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THE POLYCHAETE WORMS
DEFINITIONS AND KEYS TO THE ORDERS, FAMILIES AND GENERA

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THE
POLYCHAETE WORMS
Definitions and Keys to the
Orders, Families and Genera
By Kristian Fauchald
NATURAL HISTORY MUSEUM

OF LOS ANGELES COUNTY
In Conjunction With
THE ALLAN HANCOCK FOUNDATION
UNIVERSITY OF SOUTHERN CALIFORNIA
Science Series 28
February 3, 1977
ti
TABLE OF CONTENTS
PREFACE VII
ACKNOWLEDGMENTS IS
INTRODUCTION 1
CHARACTERS USED TO DEFINE HIGHER TAXA 2
CLASSIFICATION OF POLYCHAETES 7
ORDERS OF POLYCHAETES 9
KEY TO FAMILIES 9
ORDER ORBINIIDA 14
ORDER CTENODRILIDA 19
ORDER PSAMMODRILIDA 20
ORDER COSSURIDA 21
ORDER SPIONIDA 21
ORDER CAPITELLIDA 31
ORDER OPHELIIDA 41
ORDER PHYLLODOCIDA 45
ORDER AMPHINOMIDA 1 00
ORDER SPINTHERIDA 1 03
ORDER EUNICIDA 1 04
ORDER STERNASPIDA 114
ORDER OWENIIDA 114
ORDER FLABELLIGERIDA 1 15
ORDER FAUVELIOPSIDA 117
ORDER TEREBELLIDA 118
ORDER SABELLIDA 135
FIVE "ARCHIANNELIDAN" FAMILIES 152
GLOSSARY 1 56
LITERATURE CITED 1 61
INDEX 180
Preface
HE STUDY of polychaetes used to be a leisurely I apologize to my fellow polychaete workers for
T occupation, practised calmly and slowly, and

the presence of these worms hardly ever pene-
trated the consciousness of any but the small group
introducing a complex superstructure in a group which
so far has been remarkably innocent of such frills. A
great number of very sound partial schemes have been
of invertebrate zoologists and phylogeneticists inter- suggested from time to time. These have been only
ested in annulated creatures. This is hardly the case partially considered. The discussion is complex enough
any longer. without the inclusion of speculations as to how each
Studies of marine benthos have demonstrated that author would have completed his or her scheme, pro-
these animals may be wholly dominant both in num- vided that he or she had had the evidence and inclina-
bers of species and in numbers of specimens. In some tion to do so.
areas they are even dominant in biomass. If any state-
ments are to be made about the biology of the benthic Kristian Fauchald
environments, some note must be taken of the poly-
chaetes. Furthermore, the keeping of some of these 1 9 May 1976
animals in culture has proved feasible, and some Los Angeles
polychaetes even have become famous for their value
as test animals in polluted areas. The development of
some polychaetes appears to be genetically interest-
i ng, and studies of evolutionary rates and genotypic
and phenotypic adaptations in these worms may prove
very illuminating to our understanding of the benthic
environment.
All of these developments make it important that
the major morphological and anatomical features be
well understood, and preferably that the terminology
and taxonomic categories be agreed upon by most
workers. This review is an attempt at summarizing
current information about the taxonomy and morphol-
ogy of these animals. During the reviewing process,
it became painfully obvious to me how exceedingly
poorly known the group is and how few generaliza-
tions can be made on the ecology and evolution of
the polychaetes. I hope this review will bring forth a
spate of investigations, by persons wishing to prove
me wrong in my phylogenetic speculations, but also
by persons willing to put in the hard work needed to
fill in the gaps in our knowledge.
Acknowledgments
HIS REVIEW is a direct outcome of a lecture I
T gave during a class in polychaete biology at

Santa Catalina Marine Biological Laboratory in
1 973. It has benefited greatly from discussions with
all of the students in the class, but perhaps especially Bill
Kennedy was instrumental in pushing me into giving the
l ecture in the first place. I have also discussed parts of the
paper with Ray Emerson, Tom Kauwling, Fred Piltz and
Bob Smith. My co-teacher at Catalina, Paul C. Schroe-
der, was a great help in formulating some of these ideas.
Some of the ideas presented here were also discussed
with the late Dr. Olga Hartman, whose sound advice was
always valued by me. Dr. Pat Hutchings advised me of
some errors in the terebellid key, for which I am grate-
ful. The good illustrations were made by Ms. Catherine
Link; the others I am responsible for. Ms. JoAnne
Woodcock expertly typed most of the manuscript,
prepared the index and cleaned up my English where
needed, for which I am grateful. I am also very grateful
to Mr. R. Edward Ostermeyer for seeing the paper
through.
I would wish to thank Dr. Robert J. Lavenberg,
Natural History Museum of Los Angeles County, for
prodding me into writing this book in this format and for
his constant encouragement and enthusiasm for what
grew from a moderately long key to families to a rather
more massive offering. I am also very grateful to Dr.
Bernard C. Abbott, Allan Hancock Foundation, Uni-
versity of Southern California, for his support and en-
thusiasm and for letting me have the time to do the
writing on this paper.
THE POLYCHAETE WORMS
DEFINITIONS AND KEYS TO THE ORDERS, FAMILIES AND GENERA'
By KRISTIAN FAUCHALD'
ABSTRACT: A review of the classification of the Class Polychaeta (Annelida) with comments on the
characters used to identify the different included taxa has led to the recognition of seventeen orders. All taxa
down to the generic level are defined and a phylogenetic sequence suggested. Keys are presented to the
families and genera of the Polychaetes.
INTRODUCTION and the setae, if present, only rarely occur in bundles.

These two groups were considered more advanced
Polychaeta is part of the old, diffuse concept Vermes,
than the marine, dioecious polychaetes. The poly-
a group that perhaps can best be defined as comprising
chaetes have been defined for the last seventy years
all animals that are longer than wide and non-vertebrate.
as dioecious, marine annelids with parapodia bearing
For the last seventy years or so, the polychaetes have
numerous setae. They also should have anterior appen-
been grouped with the oligochaetes and hirudineans and
dages of various sorts (antennae, palps, tentacular
a few smaller groups into the phylum Annelida. This
cirri) and the gonadal ducts should be simple.
phylum contains segmented, coelomate worms in which
These definitions work if some of the smaller groups
a secondary loss of segmentation may have taken place,
are disregarded. If these groups are taken into account,
but in which traces of such segmentation at least can be
as they must, the only separation that consistently can
recognized internally. Other, more formal definition
be made between the oligochaetes/leeches and the
of the phylum can be found in standard text-books.
polychaetes, is the presence in the former grouping
Definition of the annelid classes has varied, however,
of hermaphroditic gonads limited to a few segments.
everyday recognition of members of the three major
classes has never been a problem: The Hirudinea con- Some hermaphroditic polychaetes are known, but these
usually have gonads in a large fraction of the total num-
tains the leeches, the Oligochaeta the earthworms and
ber of segments. It is then difficult to give a good,
their allies, and the Polychaeta marine worms, such
consistent and practically useful definition of what
as the sandworm (nereids) and bloodworms (glycerids)
is meant by a polychaete, but a definition along the
used for bait in parts of the world. The three classes
lines suggested below, should separate them from the
appeared also, at least grossly, to be separated ecolog-
other annelids with reasonable accuracy.
ically in that the leeches were supposed to be parasitic,
The polychaetes are multi-segmented annelids with
the earthworms terrestrial and the polychaetes were
parapodia; setae are present in distinct fascicles. They
most common in the marine environment. This separa-
are dioecious and have simple exit ducts from the
tion is obviously unsatisfactory and as information
about these animals accumulated towards the end of
the last century, most workers settled on a grouping
which associated the leeches and the earthworms with 1 REVIEWCOMMITTEE FOR THIS SCIENCE SERIES
each other in one group opposed to the polychaetes. ROBERT J. LAVENBERG
Members of both the former classes are hermaphroditic DONALD REISCH
and have various complex glands associated with repro- ' Allan Hancock Foundation, University of Southern California,
duction. Furthermore, both leeches and earthworms Los Angeles, California, 90007. Contribution Number 358 of
lack parapodia (fleshy unjointed segmental appendages) the Allan Hancock Foundation.

2 NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY Science Series 28
gonads. They are usually marine, more rarely fresh- 1 962, pp. 424-425; Clark 1969, p. 47). Polychaete
water and only rarely terrestrial or parasitic in habitat. taxonomists have tended to disregard these attempts
Any of these features need not be present and none of and have continued to treat the polychaetes as if the
them is essential for the recognition of an animal as class consisted of two orders (Fauvel 1958, pp. 166-
a polychaete. 1 90; Hartmann-Schroder 1971, p. 29) or subclasses
This topic has been treated in considerably greater (Uschakov 1955a), or have treated the group as if it
detail by Clark (1969) and to a lesser extent by Fau- consisted of about 75 distinct and unrelated families
chald (1974a). (Hartman 1968, 1969). The problem with all proposed
A key morphological feature and at the same time schemes is that they are internally inconsistent. Fur-
one of the most important taxonomic characters of the thermore, they give no better solutions to classificatory
polychaetes is the setal (chaetal) construction. The problems than the old, admittedly artificial, separation
setae are ectodermal derivatives, formed by ectodermal into two orders.
cells that during the development have migrated to a The three most ambitious recent proposals were by
position well below the rest of the ectodermally derived Dales (1962), Storch (1968) and Clark (1969). Dales
epidermis. Each seta consists of a bundle of filaments used the variable structures of the eversible stomodeal
laid down by a basal chaetoblast and up to several region (pharynx) to separate different groups. The
lateral cells. The material in the setae is a glycoprotein, arrangement of the body-wall musculature was used
consisting of chitin (a polysaccharide) and a protein by Storch. Clark used a variety of different structures
cross-linked at the time of formation. The formation to characterize his eight orders. These authors gave
of structural details in the setae is very well controlled, no formal definition of any taxon above the family
but exactly how this takes place has only partially been level (except by inference from contained taxa) and
clarified. The best current review of this topic was it has been difficult to evaluate their schemes.
made by O'Clair and Cloney (1974) from which most
of the above information has been gleaned.
Polychaetes traditionally are separated into two large
orders, ERRANTIA and SEDENTARIA (Audouin and CHARACTERS USED TO DEFINE
Milne Edwards 1834, pp. 24-26). The separation is HIGHER TAXA
based on the development of the anterior end and the
life habits of the included species. Major anatomical and morphological features were
The errants are supposed to have a large number of reviewed during a study of the phylogenesis of the
equal body-segments. The anterior appendages are few polychaetes (Fauchald 1974a). Below is given a survey
in number and differentiated into palps, antennae, of the findings with an expanded discussion of their
tentacular cirri, etc. These worms are considered free- taxonomic aspects.
living and, generally, should be rapacious in habits. A. Prostomium. The prostomium usually is dis-
All polychaetes with jaws are included in this order; tinct and may have or lack appendages. In several
thus the onuphids, despite their tubicolous habits, are families it is more or less fused with the peristomium
considered errants since their large jaw-apparatus and the first segments. The degree of fusion is difficult
resembles the jaw-apparatus in other, non-tubicolous to determine even in an examination of the nervous
eunicidlike animals. system so the degree of distinctness of the prostomium
The sedentaries are supposed to have a limited num- is a character that can have no great taxonomic value
ber of body segments. The body may be separated into (see Benham 1894, 1896). Prostomial appendages in-
different regions. Anterior appendages may be absent clude antennae and palps. Antennae are innervated
or a few to many similar appendages may be present. through single roots directly from the brain; palps
The sedentaries have short parapodia associated with always have double roots, either from the brain or from
their tubicolous or burrowing habits and are usually the circumesophageal ring (Akesson 1963; Orrhage
deposit- or filter-feeders. 1966). Antennae are always sensory; palps may be
These definitions have not changed much over time sensory or may be used as feeding appendages. The
(cf. Grube 1850, p. 281 and tables; Fauvel 1923a, presence of either one or both categories of appendages
pp. 27-29; Hartmann-Schroder 1971, p. 29). The ad- is considered here of great importance. The position
vantage of the system is that the bulk of the 8,000+ of the palps varies from ventral to dorsal, from frontal
described species of polychaetes separates into two to occipital. The position and function of the palps fur-
roughly similar groups in terms of numbers of species nish important taxonomic characters. It is impossible
and genera as well as families. The separation is other- to distinguish any other classes of prostomial appen-
wise unsatisfactory since neither order can clearly be dages either on morphological or anatomical grounds.
defined. Several attempts have been made to subdivide B. Peristomium. The larval peristomium is the
the polychaetes in a more acceptable manner (Dales i mmediate prototrochal region of the trochophora larva;

1976 THE POLYCHAETE WORMS 3
it may persist as the adult peristomium at least in some a constant number, or at least only a few alternative
forms and it appears to be completely pre-segmental in numbers (nereids, syllids, phyllodocids, etc.). Fused
nature, at least in some forms (Akesson 1967). How- segments are often present, even if tentacular cirri
ever, the structure called the peristomium in most are absent.
taxonomic studies consists of a fusion of this larvally It would be valuable to distinguish the two kinds of
derived structure and one or more true segments. The peristomia, but current usage seems ingrained and little
l arvally derived peristomium may carry a single pair would be gained by coining a new term; it should,
of dorsal cirri called peristomial cirri. The fused seg- however, be remembered that the current term may
ments may carry parapodial remnants called tentacular cover two very different structures.
cirri. The number of tentacular cirri vary from one to C. Eversible pharynx. Most polychaetes can everse
four pairs in the hesionids; other families tend to have a part of the anterior digestive tract. Two different
constructions can be recognized (Dales 1962). A ventral
plate-muscle pharynx is present in several forms (Eu-
nicea, Amphinomida, etc.); others have an axial phar-
ynx that is developed symmetrically, or at least nearly
symmetrically. The axial pharynx may be followed by
a strongly muscular region (nereids, nephtyids, glyc-
PALP erids, etc.) or this musculature may be absent (arenic-
ANTENNA olids, maldanids, etc.). Usually each family has a
PROSTOMIUM characteristic kind of pharynx, but in some families (e.g.,
PRESEGMENTAL_ PERISTOMIAL CIRRUS Spionidae) both plate-muscle and axial pharynges are
T PERISTOMIUM
SEGMENT Iosotlg.rousl
present (Orrhage 1966). Some plate-muscle pharynges
are poorly muscularized and may be difficult to dis-
SETIGER
tinguish from weakly developed axial pharynges so the
PARAPODIUM apparent overlap in distribution of the two kinds of
SETA pharynges may in part be due to definitory problems.
This problem suggests that the structure of the pharynx
cannot be used as the single definitory character for
higher taxa. The detailed structure of the pharynx,
especially the equipment of jaws, teeth and other
chitinized structures associated with the anterior end,
are very important characters at the generic and specific
levels. The variability of the jaw-apparatus of several
members of the super-family Eunicea is presently
under investigation. Preliminarily, it appears that the
detailed structure of the jaws is correlated very pre-
POSTSECMENTAL PYGIDIUM
ANUS cisely to other variable morphological features and
ANAL CIRRUS to environmental variables (Fauchald and Smith, in
preparation).
D. Parapodia. Polychaete parapodia can be bi-
ramous, with both noto- and neuropodia developed,
or uniramous with only the neuropodia developed. In
the latter case, the notopodia are considered secondarily
reduced (Fauchald 1974a). The presence of notopodia
is a very important character at the supra-familial and
ordinal levels as is the presence of acicula and setae.
The detailed development of each ramus with the vari-
ous parapodial lobes and cirri is very i mportant at the
NOTOPODIUM generic and specific levels. The presence of branchiae
associated with the parapodia is of variable importance.
NEUROPODIUM
The presence of branchiae may not even be considered
VENTRAL NERVE PARAPODIUM
a specific character (Fauchald 1970 on Eunice (Nicidion)
cariboea and Palola spp.); in other cases the presence
may be used as a generic character (Asychis and Bran-
FIGURE 1. Diagrams showing the major morphological features chioasychis). Generally, however, the presence of
of a generalized polychaete. branchiae is a specific character, but with generic

importance where warranted by evidence. Branchiae tend to be rather poorly treated. The most complete,
are of sufficient biological importance, so fairly good and thus most easily identifiable specimens will be
evidence must be presented to demonstrate that these caught with gear that takes chunks of the environment
structures are of less than specific importance. and in situations where the animals are allowed to
E. Setae. Numerous kinds of setae have been de- crawl out, or where the material is gently screened.
scribed (uncini, limbate setae, pectinate setae, sub- Even in S.C.U.B.A. collection it is advisable to col-
acicular hooks, composite spinigers and falcigers, etc.). lect masses of material, rocks, seaweed tufts, sand-
The basic kind of setae in each parapodial ramus is samples, etc. and put them in separate plastic bags
usually a family character, but not uniquely so. Thus, or similar waterproof sample bags for later treatment.
all phyllodocids have composite setae, but not all Most polychaetes are small and active animals; it is
polychaetes with composite setae are phyllodocids. The therefore useless to put a bottom sample in the ordi-
detailed construction of the setae is important at the nary game-bags used in diving, since most of the worms
specific level. The importance of accurate examination will escape before the sample can be treated further.
of the setae still is underestimated by most taxonomists; Shipboard sampling can be done with any of several
precise observations require close microscopic work kinds of a series of gear, quantitative as well as qual-
to elucidate them and this kind of work may be neces- itative. If dredges are used, I recommend hauls as short
sary even in routine identifications. Moreover, recent as practicable, since the churning of the material in
studies with scanning electron microscope have demon- the dredge will tend to grind up the polychaetes or
strated clearly the importance of accurate work on disturb them enough to make them autotomize appen-
the setae (Thomassin and Picard 1972). dages and, often, the whole posterior end.
F. Nephridia. The structure and distribution of Samples of hard substrates, algal mats, rubble, etc.
nephridia have been used at the subfamilial and generic should be put in a large container of some sort, covered
levels (Hessle 1917 on terebellids). The character may completely with sea water and be left standing undis-
be of wider usefulness at higher levels, but the varia- turbed for several hours. The samples should be placed
tion in these features has been too little studied in in the dark or at least in the shade and should be kept
most families to make the character useful at the pres- cool, though not necessarily refrigerated below the
ent time. ambient temperature at the sampling site. It is espe-
Most polychaete families are characterized by a cially important to leave the sample in the dark if it
complex set of features and cannot be identified by contains algae in quantity. As the oxygen concentra-
reference to a single structure. The proposed taxonomic tion in the water decreases, the polychaetes will leave
schemes have failed because they did not take this the substrate and congregate around the rim of the
into account. They all were based on two or perhaps vessel at the air-water interface. They can easily be
three manifestations in one important structure. By scooped from the surface with either a small screen
defining each manifestation precisely, apparent "inter- (0.5 mm mesh-size) or simply a spoon. If the sample
mediate" forms appeared and had to be included in contains a large number of motile, large animals,
one or another category as an exception or be left out- such as crabs or brittle-stars, it is best to remove most
side the proposed system. Usually, the "intermediate" or all of these as soon as possible.
form would be loosely appended to one taxon or an- The samples should be left standing for several
other; the definition of the taxon would then be left hours (2-12 hours) depending on the lighting, the
intact. It became impossible to find these "hidden" quantity of water in relation to sample size, the tem-
taxa in the system. perature, etc. After the samples have been treated in
this manner, the substrate should be carefully sorted
through-plucked to pieces if necessary-and the
SOME USEFUL TECHNIQUES water screened for animals that left the substrate but
did not reach the surface. This whole process should
The following comments are necessarily incomplete be done as quickly as possible, since polychaetes de-
and are meant only as a first-hand guide to work on teriorate very rapidly after death. The method will not
polychaetes, especially the handling necessary to per- quantitatively remove polychaetes, since some poly-
form identifIcatory work, excluding the handling neces- chaetes are unable to leave their tubes or burrows and
sary for other kinds of investigations on either live others generally do not approach the surface, even
or dead polychaetes. The main topics covered are when the water becomes very foul. The process can
collection, screening, fixation and preservation and be speeded up by adding 7% MgCl,, but we have not
the most common techniques in laboratory handling. found this to be any great advantage.
Collection. -Polychaetes are found nearly every- Screening.-Samples of soft substrates, such as
where in the marine environment and thus can be sands and muds, must be screened. Quantitative benthic
caught with every kind of gear imaginable. They are studies now routinely use t mm screens, but these un-
soft-bodied animals, and hand-collected specimens dersample the polychaetes badly, both in numbers of

species and in specimens. The use of 0.75 mm screens water as possible. The pan is then flooded with a solu-
does not improve matters greatly, but 0.5 mm screens tion of 7% MgCl, in sea water and allowed to stand
appear to catch most polychaetes quantitatively. In for about half an hour. Then the contents of the pan
inshore areas, the sands and gravel will make the use are poured carefully through a screen and transferred,
of 0.5 mm screens rather impractical, but if a complete with as little water as possible, to a sample jar. The
survey of the fauna is contemplated, this cannot be narcotizing solution can be used several times if so
avoided. Thus, the objective of the study undertaken desired.
must be considered carefully. I would generally recom- Standard fixating agent for polychaetes is 10% neu-
mend that, for each area studied, at least some samples tralized formalin in sea water. The most commonly
be screened with a double set of screens, both I .0 and used neutralizing agent is borax (Na,B 4 O7, technical
0.5 mm, and the results compared so that the level of grade). The sample should fill no more than one-third
inaccuracy engendered by the use of 1.0 mm screens of the sample jar and the jar should be completely
can be estimated. This has to be done for every major filled with the formalin solution. The jar should be
sediment type and cannot be guessed at from one sedi- capped and gently but thoroughly inverted several
ment type to the next, nor from one geographical loca- times to get complete mixing. Allow the sample to
tion to the next. This is because the fraction of small settle, decant off about one-half of the solution and
species appears to vary geographically. fill the jar with fresh formalin solution. It is much
Samples should under any circumstances be washed better to split a sample into several jars than to fill
gently with very large quantities of water. It is useful one jar completely with the entire sample and get
to have the water prescreened so that pelagic organisms incomplete and unsatisfactory fixation.
can be avoided, and deep-water samples should be Histological fixatives can also be used on bulk
treated with water as cold as possible. Nothing is samples, and in general such fixation is better on small,
gained by hurrying the screening process, but each fragile polychaetes if done prior to any sorting. If
sample should be screened as soon as possible after such fixatives are used, the ratio between sample and
getting it on deck. Deep-water samples that cannot be fixative must be even lower than the one indicated
screened immediately should be refrigerated. It is of above, and the fixative should be changed twice in
the utmost importance for good results that the screen- rapid succession to avoid dilution effects.
ing process be done carefully; poorly screened samples Samples should be left in formalin for at least 24
contain a large number of mangled specimens, and hours and can be left in the fixative for several weeks.
such specimens usually cannot be identified. The net However, after 24 hours, the samples are ready to be
result will be a waste of sampling time and effort and, transferred to the preservative, usually 70% isopropyl
not least, loss of time needed for identification of the or ethyl alcohol in distilled water. Before transfer,
specimens after the samples have been returned to the the samples must be washed in fresh water once or
laboratory. In terms of the time and effort needed for twice to remove the salt; if this is not done, setae and
the different parts of the processing of a single sample, other details will become the crystallization sites for
we now generally calculate that on the average it takes salt crystals, and these are difficult to remove after
about 50 minutes of shipboard time to take and process they have formed. One change of preservative is neces-
a single shelf sample with a box-core and about three s ary to ensure full strength.
weeks of manhours to adequately treat and interpret DO NOT ATTEMPT TO USE ETHYL ALCOHOL
this sample in the lab. Thus, the few minutes of ship AS A FIXATIVE, even if sensitivity to formalin be-
time to be gained in preparing the samples poorly will comes a problem. The specimens become completely
be offset by the need for a larger number of replicate unusable after a short period of time, and again the
samples to get adequate numbers for mathematical fraction of unidentifiable specimens goes up drastically.
treatments in the lab. Postfixation with formalin of material originally fixed
Once the sediment has been removed from the in alcohol does not work.
screens as completely as possible, the retained material Specimens treated as recommended above usually
should be chased down to one side of the screen with retain most of their appendages, and a large fraction
the help of a gentle stream of water applied to the of complete specimens is usually present. However,
outside of the screen. Do not sort or pick through the a certain number of incomplete specimens are to be
material on the screens: non-preserved polychaetes expected in any treatment, especially with standard
should be handled as little as possible and never moved narcotizing time as suggested above. The treatment
at all except with the help of a gentle stream of water suggested is rather more elaborate than usual, but we
applied to the outside of screens or elsewhere. have found that we are amply rewarded by a much
Fixation and Preservation. -For the last few years higher than usual fraction of identifiable specimens.
we have routinely narcotized the whole sample as Thus, less replication of samples is needed in benthic
retained on the screens. The material on the screens is surveys and this in itself represents a savings in both
washed down onto a large enamel pan with as little sea time and money.

Laboratory Treatment of Samples, Identificatory siderably more efficient to identify the polychaetes
Techniques. -The equipment needed to identify poly- family by family in larger numbers. It is much easier
chaetes includes one stereo microscope per person and to compare specimens from different samples, and the
one compound microscope per two persons, as the number of dubious identifications can be decreased
minimum. The compound microscope must be capable drastically by this means. If the samples are to be
of magnifications up to 1100 times, and the stereo stored as units, the vials can always be reunited after-
microscopes should have magnifications to about 100 wards if desired.
times. A focusable microscope lamp is necessary for When polychaetes are being identified, certain
use with the stereo microscope. Each person will also standard observations should always be made. Always
need two pairs of watchmaker's forceps, two needles find the anterior end and take note of the number and
(we use insect pins glued to applicator sticks with arrangement of anterior appendages. Scan the body for
epoxy glue), fine scissors, a small scalpel (we use obvious differences in parapodial structures and for
pieces of razor blades glued to applicator sticks with such features as the position and number of branchiae.
epoxy glue), a small bottle of glycerol alcohol mixed Most specimens preserved as indicated above will be
with one-half glycerol and one-half 70% alcohol, a relatively easy to handle with two pairs of forceps,
bottle of immersion oil, depression slides and flat and it is usually much easier to move the specimen
slides, cover slips, petri dishes of various sizes (pref- than to move the dish. This sort of scan is usually all
erably with tight-fitting lids), cotton, paper, and #2 that is needed to get a polychaete to family.
pencils (HB or F works well). To identify the animals to genus and species, more
The sorting and identification of polychaetes is a accurate observations are usually necessary and various
two-step operation. Each sample should first be sorted dissections must be performed. It is always necessary
to family under a stereo microscope. The samples to remove a parapodium, if nothing else, because a
must be sorted while completely submerged in alcohol, good look at the setae is needed. This can be done
and since light from the microscope lamp is apt to with the use of two pairs of forceps, a scalpel or iris
evaporate the alcohol it should be refreshed from time scissors. For most medium-sized polychaetes in good
to time. Polychaetes always have to be treated while condition, just pulling off a parapodium with a pair
completely submerged, and after being transferred to of forceps is the easiest method. Care must be taken
alcohol cannot be left dry for more than a few minutes at that both parapodial rami and associated cirri and
a time. Small polychaetes tend to dry out very quickly, branchiae come off. Some workers find it easier to
and dried out polychaetes are largely unidentifiable. use scissors or scalpel, and these instruments must
Sorting can be done directly into vials completely be used on poorly preserved specimens or on larger
filled with alcohol, and each vial receives a label with animals. The parapodium should be mounted on a
the name of the contained taxon, as well as with the slide; larger ones must be mounted on depression
station number for that particular sample. It is impera- slides, but normally a flat slide is better, since depres-
tive to do this immediately rather than having to re- sion slides cannot be used with high-power compound
member the content and position of each vial later. microscopes. Generally, the parapodium should be
Use good, high-rag-content paper of sufficient weight mounted with the anterior side facing the observer;
(20 pound or higher) and a good pencil. The labels however, in certain families and genera, a posterior
should be big enough to stand up in the vial unsup- view may be more informative. Before mounting the
ported, but not so large as to cover the contents, and parapodium, look at some appropriate parapodial
they should end up well below the upper margin of illustrations showing the features to be observed. A
the vial. We prefer to use straight-sided vials, capped parapodium mounted in a dorso-ventral position gives
with cotton plugs, and to store these in jars filled with no more information than does looking at the whole
alcohol. Screw-capped vials almost invariably have a animal, so a mount showing both notopodium and
shoulder which makes it difficult to remove specimens neuropodium is necessary. We use glycerol-alcohol
and labels when needed. Screwcaps normally are not for these mounts and only rarely make permanent
air tight, so the vials will have to be stored within a mounts.
l arger jar regardless. Which parapodium should be removed depends on
After the sample has been sorted to family, we col- which family is being studied. Generally, a median
lect the members of each family in a single jar. Thus, parapodium from a long series of similar-looking
the sample is no longer intact as such, but has been parapodia will be best, but in special cases the anterior-
distributed taxonomically among families. It is thus most or one specific parapodium will have to be re-
of very great importance that careful and accurate moved to study some specific detail of importance in
notes be taken on the numbers of vials for each station, that taxon. Thus, in members of the genus Pista, for
so that later the station can be reconstructed accurately example, anterior, median and posterior thoracic para-
on paper. This is done because we have found it con- podia must be removed, and in the genus Magelona

the ninth parapodium must be detailed. Again, it is made, etc. The jaws in these forms are always ob-
worthwhile to look through the key to be used before served from the dorsal side and the jaw formulae are
deciding which parapodium and, in some cases, how given from the posterior to the anterior end, the left
many parapodia, should be removed. It is also worth- jaw being mentioned first in each formula.
while to scan the body of the animal carefully before The pharyngeal lining has characteristic structures
deciding on a specific parapodium, to see that the in members of several families; it is usually not neces-
setae are as complete as possible so no extra mounts sary to remove the lining in nephtyids and phyllodocids,
of setae become necessary. Setal distribution varies but in order to characterize the lining of members of
from one group of polychaetes to another; if it is pos- glycerids and goniadids, this must be done. It is of
sible to remove one single parapodium and get all setal great importance that the lining be well oriented in
types represented at once, this saves wear and tear on the goniadids, since the position of the different kinds
the specimens and also saves time in preparation. of pharyngeal organs is considered of taxonomic im-
Generally, parapodial structures are most easily portance. The fine structure of the glycerid pharyngeal
observed at relatively low magnifications under the organs cannot be seen except in oil immersion, and
compound microscope, but critical decisions, especially critical lighting is of the utmost importance for a
on structures on top of a thick preparation, may be clarification of these structures.
most easily followed at higher magnifications. Setal Parts that have been dissected out are best put in
structures, especially the presence of fine hairs along a small, separate vial stored within the larger vial
the cutting edge of simple setae, are best seen under with the specimens. Leaving small parts loose in the
oil immersion with 100x objectives. It does not help vial will invariably lead to their loss.
much to use very high oculars; generally, a lOx ocular
is more than adequate. If available light is insufficient
for using the immersion objective, attempt to put im- CLASSIFICATION OF POLYCHAETES
mersion oil also between the condensor and the lower
side of the slide. Be sure to adjust the lighting of the The scheme proposed below left. also Table 1), is
compound microscope every time the magnification based on phylogenetic ideas presented elsewhere
is switched; it gives a much more satisfactory picture (Fauchald 1974a). The sequence of families indicates
of the structure and will also, in the long run, save an increasing morphological distance from the ancestral
the eyesight of the worker. polychaete as this was defined on that occasion, but
Dissection of the anterior end of the polychaete may since the several orders and families are considered
also be necessary for observation of the structure of the results of a rapid radiation in Pre-Cambrian to
the eversible pharynx or jaw structures. Important Cambrian times (Fauchald 1974a), this sequential
structures are situated in the midline, dorsally or ven- arrangement can only poorly represent the phylogenetic
trally or both; thus, a dissection should avoid cuts pattern.
into the midlines of the animals. We generally make Major anatomical as well as morphological features
a longitudinal slit on the dorsal side, well lateral to were used to define the orders. This may make them
the midline. The length and position of the slit will difficult to use in practical taxonomic work. The frame-
depend on the purpose of the dissection and on the work formed by including the anatomical features is
relative position of the eversible pharynx. The pharynx more satisfactory in that each order now can be defined
may be preserved in the completely retracted position to exclude all non-members.
or in various stages of eversion; jaw structures are Theoretically different evidence should be used
usually situated anterior in the body, even in forms to define each taxonomic level. One feature (e.g., the
with a long pharynx. If the purpose of the dissection structure of the eversible pharynx) once used at one
is observation of the lining of the pharynx in one of taxonomic level, should not be used at another (lower)
the forms with a long pharynx, the slit can be made level within the same classificatory plan. This sort
farther back than usual. of separation was attempted here, but was only partially
We usually continue by making transverse slits at successful, in that one feature (e.g., the structure of
both ends of the longitudinal one. This makes it pos- the eversible pharynx) may have been used at the
sible to lift a flap of tissue containing the body-wall family level in one order, but at the sub-ordinal or
proper. We have found it most useful to leave the superfamilial levels in other orders.
pharynx and the jaws in situ as much as possible; Suborders have been recognized only where war-
they are almost invariably lost if removed. Especially ranted; no attempts were made to create intermediate
in the case of the complex jaw structures of the eunicean categories in all orders. Families not included in any
polychaetes, it is important to treat them in a similar intermediate category are listed alphabetically at the
manner in all specimens, so that they are all flattened end of each order. The sequence of families otherwise
to a similar degree when the observations are being indicates phylogenetic relationships within each order.

TABLE I
Survey of Polychaete Orders, Suborders, Superfamilies and Families.
l. O. ORBINIIDA 7. O. OPHELIIDA SO. Glyceriformia 12. O. STERNASPIDA

F. Orbiniidae F. Opheliidae F. Glyceridae F. Stemaspidae
F. Paraonidae F. Scalibregmidae F. Goniadidae 13. O. OWENIIDA
F. Questidae 8. O. PHYLLODOCIDA F. Lacydoniidae F. Oweniidae
2. O. CTENODRILIDA SO. Phyllodociformia Suborder not 14. O. FLABELLIGERtDA
F. Ctenodrilidae F. Phyllodocidae recognized: F. Flabelligeridae
F. Parergodrilidae F. Alciopidae F. Iospilidae F. Poeobiidae
3. O. PSAMMODRILIDA F. Lopadorhynchidae F. Nephtyidae 15. O. FAUVELIOPGIDA
F. Psammodrilidae F. Pontodoridae F. Sphaerodoridae F. Fauveliopsidae
4. O. COSSURIDA SO. Aphroditiformia F. Tomoptendae 16. O. TEREBELLIDA
F. Cossuridae SF. Aphroditacea F. Typhloscolecidae F. Sabellariidae
5. O. SPIONIDA F. Aphroditidae 9. O. AMPHINOMIDA F. Pecunarudae
SO. Spioniformia F. Polynoidae F. Amphinomidae F. Ampharetidae
F. Apistobranchidae F. Polyodontidae F. Euphrosinidae F. Terebellidae
F. Spionidae F. Pholoididae 10. O. SPINTHERIDA F. Trichobranchidae
F. Magelonidae F. Eulepethidae F. Spintheridae F. Bogueidae
F. Trochochaetidae F. Sigalionidae 11. O. EUNICIDA 17. O. SABELLIDA
F. Poecilochaetidae SF. Chrysopetalacea SF. Eunicea F. Sabellidae
F. Heterospionidae F. Chrysopetalidae F. Onuphidae F. Sabellongidae
SO. Chaetopteriformia F. Palmyridae F. Eunicidae F. Caobangidae
F. Chaetopteridae SF. Pisionacea F. Lumbrineridae F. Serpulidae
SO. Cirratulifonnia F. Pisionidae F. Iphitimidae F. Spirorbidae
F. Cirratulidae SO. Nereidiformia F. Arabellidae Families of uncertain
F. Acrocirridae F. Hesionidae F. Lysaretidae affinities:
6. O. CAPITELLIDA F. Pilargiidae F. Dorvilleidae F. Dinophilidae
F. Capitellidae F. Syllidae Super-family not F. Nerillidae
F. Arenicolidae F. Calamyzidae recognized: F. Polygordiidae
F. Maldanidae F. Nereidae F. Histriobdellidae F. Protodrilidae
F. Antonbmunidae F. Ichthytomidae F. Saccocirridae
All known taxa to the generic level have been de- Because of this and because of the general variability
fined. Some taxa can be characterized by a single of key features within each family, several families
unique structure. The absence of this structure in all key out at different points.
other taxa at that level has been left unstated to save The generic keys are dependent on the presence of
space. deciduous features. Most of these keys are short enough
Preferably, keys should have been made first to to allow scanning of the total key for necessary cor-
order and within each order, to family. For several rections if such features have been lost.
reasons, this approach was abandoned. Identification Under all circumstances, identifications made through
of orders may be possible only after a detailed ana- the key should be checked by using the definitions
tomical study; the orders are justifiable scientifically, of genera given. Introduction of a genus into the faunal
but rather difficult in practical taxonomic work, as lists should not be attempted based on identifications
mentioned above. The general approach to keys taken made through the keys in this paper. For publication
here is that they are tools to make it possible to identify purposes, the original literature must be consulted.
the taxa swiftly and easily and with as little damage A name erroneously introduced in the literature for
to the specimens as possible. For that mason one master any area is in practice indelible. Sloppiness has been
key to families is proposed. Furthermore, all keys the cause of mom errors in the polychaete literature
were made reversible. If one has a fairly good idea of than all other causes combined.
what kind of animal one has at hand from an illustration The keys and definitions given are wholly inadequate
or otherwise, one can work backwards into the key to as a base for description of new genera, especially in
check the identification. The keys were made strictly the larger families. The keys are intended as aids in
dichotomous; trick wording was avoided. However, as getting a first approximation in identification. Once
in all other keys, the usage of adjectives and adverbs achieved, a series of very valuable regional handbooks
is rather different from common English usage. The are available. Such include Uschakov (1955a and
family key was intended to discriminate between speci- 1972), Hartmann-Schrdder 1971), Hartman (1968,
mens that had lost deciduous features such as branchiae 1969), Imajima and Hartman (1964), Fauvel (1953),
or antennae, etc., in addition to complete specimens. Day (1967) and Banse and Hobson (1974). Older, but

still indispensable are the handbooks by Fauvel (1923a, will be easier to debate the classification and phylogeny
1 927), Friedrich (1938) and Berkeley and Berkeley of polychaetes sensibly; as stated by Clark (1969), the
(1948, 1952). These books can be used for a second polychaetes do present an intractable problem of phy-
approximation to the identificatory problems. How- logeny for the time being.
ever, before publication, the original research papers
must be consulted. ORDERS OF POLYCHAETES
The review below is separated into several distinct
parts. The family key is followed by an order by order The following polychaete orders are recognized:
review of all the families. For each family, a definition ORBINIIDA, CTENODRILIDA, PSAMMODRILIDA,
may be followed by a brief note, especially noting COSSURIDA, SPIONIDA, CAPITELLIDA, OPHE-
features useful in field identification. Major recent LHDA, PHYLLODOCIDA, AMPHINOMIDA, SPIN-
reviews are also mentioned at this point. Then comes THERIDA, EUNICIDA, STERNASPIDA, OWENI DA,
a key to genera and definitions of all contained genera; FLABELLIGERIDA, FAUVELIOPGIDA, TEREBEL-
followed by taxonomic notes. These explain new taxo- LIDA, SABELLIDA.
nomic combinations and taxa and specific positions Members of the old order ERRANTIA are separated
taken in this paper. The type-species is named for all into three orders, PHYLLODOCIDA, by far the largest
genera and an approximate number of species is given. with most of the well-known families, AMPHINO-
Finally, a list of invalid genera has been added. MIDA and EUNICIDA. In addition, the small ecto-
Illustrations are given for one member of each parasitic spintherids have been assigned to their own
family: as much as possible, identification features order.
have been illustrated. The bulk of the recognized orders thus comes from
A glossary contains most of the terms used, except the old collective group (or order) SEDENTARIA.
those in general usage in invertebrate zoology; where This group was never adequately defined, because two
necessary, small line drawings have been added to very disparate subgroups had to be included; the highly
the glossary to illustrate idiomatic usage, etc. modified species now included in the orders STERN-
The literature cited contains references to the original ASPIDA, OWENIIDA, FLABELLIGERIDA, TERE-
descriptions of all genera listed, except the invalid BELLIDA and SABELLIDA, and the simple-bodied
ones and to major revisions and handbooks. forms now included in the orders ORBINIIDA,
It should be noted that this paper contains little that CTENODRILIDA, PSAMMODRILIDA, COSSURIDA,
is wholly new; in most cases I have followed the most SPIONIDA, CAPITELLIDA and OPHELIIDA. The
recent major revision of any family, or followed clues latter seven orders contain structurally rather simple
indicated by revisions currently under way. It is hoped forms, but this should not be taken to indicate that
that this compilation of information may make it sim- the orders for that reason are related closely to each
pler to get more complete revisions made for each other. These forms are about as far apart as any other
family where needed. However, such a revision must grouping of polychaetes that might be proposed,
be based on materials, not on descriptions. A major judging from differences in tagmatization, parapodial
revision is a long and very laborious process, but must development and setal distribution. They could have
be undertaken; it is wholly unsatisfactory to base such been included under the old concept DRILOMORPHA
revisions on previous descriptions only, since inter- (cfr. Uschakov 1955a, Dales 1962; Clark 1969), but
pretation of descriptions frequently is dependent on defining this concept would have been very nearly
poorly understood and used terminology; a fact that impossible. The approach taken here, was that major
frequently has obscured close similarities in structure different body constructions were given the rank of
(cfr. Fauchald and Belman 1972; Blake 1975). order and that modifications on these major body plans
Ultimately, one would hope that by organizing and were given familial rank. Intermediate taxa were
defining each known taxon as clearly as possible, it employed only where appropriate.
KEY TO FAMILIES
Ia. External segmentation and setae absent 2

lb. External segmentation and/or setae present 4
2a (1 a). Paired antennae on the prostomium 3
2b (la). Paired antennae absent; tactile hairs along the body; small interstitial forms DINOPHILIDAE
3a (2a). Body long and slender; interstitial forms PROTODRILIDAE (in part)
3b (2a). Body short and saclike, pelagic POEOBIIDAE
4a (Ib). Body a flattened disc with indistinct segmentation; ectoparasitic SPINTHERIDAE

4b (lb). Body not a flattened disc; segmentation usually distinct, if indistinct, then body clearly longer than
wide 5
5a (4b). Dorsum with series of elytrae (scales) or distinct elytral scars present at the dorsal side of notopodial
bases in several segments; felt of matted notosetae may be present 6
5b (4b). Dorsum without elytrae, elytral scars or felt 11
6a (5a). Neuracicula distally hammer-headed EULEPETHIDAE
6b (5a). Neuracicula distally pointed 7
7a (6b). Prostomium with a single median antenna; dorsum with felt, or notosetae harpoon-shaped or held
erect over the dorsum APHRODITIDAE
7b (6b). Prostomium with one to three antennae; dorsum without felt; notosetae usually distinctly lateral in
position, never harpoon-shaped 8
8a (7b). Neurosetae composite 9
8b (7b). Neurosetae simple 10
9a (8a). All posterior segments with elytrae; prostomium with one to three antennae SIGALIONIDAE
9b (8a). Elytrae alternate with dorsal cirri along the whole length of the body; one antenna present
PHOLOIDIDAE
l0a (8b). Spinning glands present; median antenna, if present, attached near the posterior or middle of the
prostomium; notosetae absent POLYODONTIDAE
l0b (8b). Spinning glands absent; median antenna attached at the anterior margin of the prostomium; notosetae
usually present POLYNOIDAE
I I a (5b). Notopodia with expanded, golden or brassy setae that more or less cover the dorsum 12
11b (5b). Notosetae otherwise (may be absent) ' 13
12a (I Ia). Prostomium with large facial tubercle and a median antenna; notosetae in rosettes .. PALMYRIDAE
12b (11 a). Prostomium without a facial tubercle; paired lateral and a median antenna present; notosetae in trans-
verse rows CHRYSOPETALIDAE
13a (I lb). Posterior end covered ventrally by a chitinized shield STERNASPIDAE
13b (lib). Posterior end not covered by a shield 14
14a (13b). Prostomium completely retracted between the first parapodia which have three pairs of tentacular
cirri, partially supported by acicula PISIONIDAE (in part)
I 4b (13b). Prostomium not completely retracted between the first parapodia which are otherwise equipped .. 15
I5a (14b). Anterior end with one or several series of long, specialized setae either covering the retractable
anterior end or forming an operculum or a series of long protective spines (paleae) 16
15b (14b). Anterior end without exceptionally long, specialized setae (NOTE: Short, strong hooks may be
present) 20
1 6a (15a). Specialized setae long and chambered, forming a protective cage around the retractable anterior
end; body with numerous epithelial papillae FLABELLIGERIDAE (part)
16b (15a). Specialized anterior setae do not form a protective cage; anterior end not retractable; skin-papillae
few and small, or absent; setae otherwise 17
17a (16b). Specialized setae slender, distally curved, often spinous; prostomium with seven anten-
nae ONUPHIDAE (part)
17b (16b). Specialized setae stout, smooth and not distally curved; prostomium without appendages or with
numerous tentacles 18
1 8a (17b). Specialized setae in a transverse row; tube conical, usually formed of closely fitted sand-
grains PECTINARIIDAE
18b (17b). Specialized anterior setae either as a fan-shaped group of paleae on either side of the anterior end or
as an operculum to the tube 19
19a (18b). Specialized setae form paleae; anterior end with two to four pairs of branchiae . AMPHARETIDAE
(part)
19b (18b). Specialized setae form an operculum; anterior end without branchiae SABELLARIIDAE
20a (15b). Anterior end, including in part the prostomium, transformed into a tentacular crown 21
20b (15b). Anterior end not transformed into a tentacular crown (NOTE: Antennae and tentacular cirri may be
crowded near the anterior end) 25
2Ia (20a). Tube calcareous; thoracic membrane present 22
21b (20a). Tube mucoid or horny, often covered with sand-grains; thoracal membrane absent 23
22a (21 a). Tube irregularly twisted or straight, sometimes coiled near base; body symmetrical; more than four
thoracic setigers SERPULIDAE

22b (21a). Tube completely coiled; body asymmetrical; four thoracic setigers present SPIRORBIDAE
(Usually considered a sub-family of the SERPULIDAE)
23a (21b). Parapodia with uncini in one or a few distinct rows; tentacular crown with smooth or pennate
radioles 24
23b (21b). Small uncini massed in dense fields in the neuropodia only; short tentacular crown with branching
tentacles OWENIIDAE (part)
24a (23a). Digestive tract recurved with anus far anteriorly; thorax without hooks except in the first
setiger CAOBANGIIDAE
24b (23a). Digestive tract straight with far posterior or terminal anus; thorax with hooks in most
setigers SABELLIDAE
25a (20b). Setiger 4 with one or a few thick spines; some median parapodia strongly modified, usually fan-
shaped; tubes parchmentlike, or, if horny, distinctly annulated CHAETOPTERIDAE
25b (20b). Setiger 4 without thick spines (NOTE: Other setigers may have modified setae); no parapodia fan-
shaped; tubes never parchmentlike, if homy, then without annotations 26
26a (25b). Numerous tentacles on the lower side of the prostomium or on the peristomium; branchiae, if present,
li mited to a few pairs of anterior setigers 27
26b (25b). Anterior end with a limited (10 or fewer pairs, usually) number of antennae and tentacular cirri,
or without appendages 29
27a (26a). Branchiae in a transverse or oblique row or grouped in two groups on either side of the anterior
dorsum, usually digitiform and smooth, more rarely bipinnate or lamellate (NOTE: Branchiae are
often lost, but scars remain); buccal tentacles retractable; uncini with teeth in one or a few
rows AMPHARETIDAE (part)
27b (26a). Branchiae, if present, on two-three successive segments, stalked or sessile, branched or as
numerous filaments, rarely smooth; buccal tentacles non-retractable; uncini with several teeth
in one or more transverse rows above the main fang (crested) 28
28a (27b). Thoracic uncini long-handled, abdominal ones short-handled TRICHOBRANCHIDAE
28b (27b). Both thoracic and abdominal uncini short-handled; sometimes with a posterior prolongation in
thoracic uncini TEREBELLIDAE
29a (26b). Prostomium with at least one pair of antennae; peristomium usually with paired palps or tentacular
cirri 30
29b (26b). Prostomium without appendages or with a single antenna; peristomium with paired dorsal palps,
maximally two pairs of tentacular cirri or without appendages 58
30a (29a). Prostomium continued posteriorly in a caruncle; large notosetae furcate; others smooth or serrated . 31
30b (29a). Caruncle absent; furcate notosetae, if present, small, or furcate setae only kind of setae present . 32
31a (30a). Notosetae arranged in transverse rows on dorsum; branchiae shorter than setae . EUPHROSINIDAE
31b (30a). Notosetae in tufts on the notopodial lobes; branchiae conspicuous branching tufts AMPHINOMIDAE
32a (30b). Palps absent 33
32b (30b). Palps present, sometimes as ventrolateral pads on the peristomium or fused to the anterior end of the
prostomium so that the latter appear cleft, but usually free and digitate 49
33a (32a). Setae absent; acicula present only in the prolonged acicular lobes of the second segment (first seg-
ment in juveniles); otherwise absent TOMOPTERIDAE E
33b (32a). Setae or acicula or both present in most segments 34
34a (33b). Prostomium long and conical; usually annulated, with two pairs of antennae at the tip 35
34b (33b). Prostomium no more than twice as long as wide, never annulated; antennae long or short 36
35a (34a). Eversible pharynx with four jaws in a cross; parapodia either all uniramous or all
biramous GLYCERIDAE
35b (34a). Eversible pharynx with more than four jaws; parapodia uniramous anteriorly and biramous pos-
teriorly GONIADIDAE
36a (34b). Jaws present 37
36b (34b). Jaws absent 40
37a (36a). Each jaws consisting of a series of denticles in a row DORVILLEIDAE (part)
37b (36a). Each jaw consisting of a single piece 38
38a (37b). Four or five pairs of jaws LYSARETIDAE
38b (37b). A single pair of jaws present 39
39a (38b). Both composite and simple setae present, parasitic in decapod crustaceans IPHITIMIDAE
39b (38b). All setae composite, parasitic in fish ICHTHYOTOMIDAE

40a (38b). One pair of antennae; interstitial forms PROTODRILIDAE

40b (36b). More than one pair of antennae 41
41 a (40b). Eyes larger than the rest of the prostomium, with well defined lenses and pigment layers ALCIOPIDAE
41b (40b). Eyes, if present, smaller than the prostomium proper, usually small pigment cups, but lensed eyes
occur 42
42a (41b). Series of large epithelial capsules on the dorsum; two or three pairs of lateral antennae and one
median, unpaired antenna SPHAERODORIDAE
42b (4tb). Epithelial capsules absent (NOTE: Dorsal cirri of phyllodocids may be inflated, but are associated
with the parapodia rather than situated on the dorsum) 43
43a (42b). Dorsal cirri large and foliose PHYLLODOCIDAE
43b (42b). Dorsal cirri, if present, cirriform 44
44a (43b). Setae absent; with a few pairs of appendages anteriorly and posteriorly, parasitic in lob-
sters HISTRIOBDELLIDAE
44b (43b). Setae present; several pairs of appendages along the body 45
45a (44b). All setae composite, pelagic LOPADORHYNCHIDAE
45b (44b). At least some setae simple, benthic or parasitic forms 46
46a (45b). With ciliary bands on the dorsum of each segment in addition to a large ciliary organ on the pros-
tomium; minute interstitial forms NERILLIDAE
46b (45b). External clliation limited mostly to small patches in sensory and respiratory organs, small or large
forms 47
47a (46b). Five prostomial antennae, inquilines in bivalved mollusks ANTONBRUUNIDAE
47b (46b). Four antennae, free-living benthic forms 48
48a (47b). Interramal cirri between the noto- and neuropodia in most forms, all setae simple .. NEPHTYIDAE
48b (47b). Interramal cirri absent; notosetae simple, neurosetae composite LACYDONIIDAE
49a (32b). Palps bi- or multiarticulated 50
49b (32b). Palps simple, sometimes fused to the prostomium so the latter appears cleft or forming ventrolateral
pads on the peristomium 52
50a (49a). Palps multiarticulated; tentacular cirri absent DORVILLEIDAE (part)
50b (49a). Palps biarticulated; at least one pair of tentacular cirri 51
51a (50b). Pharynx with paired jaws; paragnaths or soft papillae or both on the surface of the everted pharynx
or pharynx smooth, parapodia usually biramous NEREIDAE
51b (5ob). Pharynx usually without jaws; paragnaths or pharyngeal papillae always absent; everted pharynx often
with a circlet of distal papillae or lappets, parapodia often sub-biramous or uniramous HESIONIDAE
52a (49b). Palps ventrolateral pads on the peristomium; five occipital and two frontal antennae
. . . . . . . . . . . . . ..................................................... ONUPHIDAE (part)
52b (49b). Palps either fused anteriorly to the prostomium or as free ventrolateral projections; maximally five
antennae 53
53a (52b). Palps free ventrolateral projections, sometimes fused to each other 54
53b (52b). Palps fused to the prostomium so that the latter appears anteriorly cleft 57
54a (53a). Prostomium longer than wide, with a pair of antennae at the tip PISIONIDAE (part)
54b (53a). Prostomium no longer than wide (NOTE: Fused palps may make it appear longer than it is) .... 55
55a (54b). Jaws present DORVILLEIDAE (part)
55b (54b). Jaws absent 56
56a (55b). Parapodia strongly prolonged, with long supportive acicula; anterior part of the digestive tract not
visibly separated into distinct parts, pelagic PONTODRRIDAE
56b (55b). Parapodia not strongly prolonged; acicula short; anterior part of the digestive tract with a visible
proventriculus in most species SYLLIDAE
57a (53b). Eversible pharynx, if present, unarmed PILARGIIDAE
57b (53b). Eversible pharynx with four pairs of upper and one pair of lower jaws EUNICIDAE
58a (29b). Anterior end, including both pro- and peristomium without appendages (NOTE: Appendages may
be present on some anterior setigers) 59
58b (29b). Prostomium with a single median antenna, or peristomium with paired palps or tentacular cirri or
both pro- and peristomium equipped as indicated 85
59a (58a). Paired palps on the first or one of the first postperistomial segments 60
59b (58a). Paired palps absent 61

1976 THE POLYCHAEtE WORMS 13
60a (59a). Both composite falcigers and simple setae present ACROCIRRIDAE (part)
60b (59a). All setae simple, distally curved or straight, sometimes acicular CIRRATULIDAE (part)
61a (59b). With a single mid-dorsal palp on one of the first setigers (setigers 3-6 usually) .... COSSURIDAE
61b (59b). Mid-dorsal palp absent 62
62a (61b). With series of long, slender branchial filaments and tentacular and dorsal cirri along the body (often
lost, scars remain) CIRRATULIDAE (part)
62b (61b). Branchial filaments and tentacular cirri absent or limited to a few segments 63
63a (62b). Parapodia strongly reduced so that the setae appear sessile on the body-wall; all setae simple, true
capillary setae absent 64
63b (62b). Parapodia usually well developed or at least present as low folds; setae usually of several different
kinds, including in most cases true capillary setae 68
64a (63a). Thorax with series of long dorsal cirri supported by acicula PSAMMODRILIDAE
64b (63a). Body usually not clearly regionated; apart from papillae and reduced parapodia, other appendages
absent 65
65a (64b). At least some setae with an internal structure of small chambers FLABELLIGERIDAE (part)
65b (64b). No setae chambered 66
66a (65b). With a papilla between the rami of each parapodium; setae smooth and slightly recurved
. . . . . . . . . . . . . . . . . . . . . . ........................................... FAUVELIOPSIDAE
66b (54b). Without papillae between the rami; setae otherwise 67
67a (66b). Setae in four fascicles on each segment (biramous condition) CTENODRILIDAE
67b (66b). Setae in two fascicles on each segment (uniramous condition) PARERGODRILIDAE
68a (63b). Prostomium an oblique plaque, usually bordered by a flange 69
68b (63b). Prostomium pointed, rounded or blunt 70
69a (68a). Setae include anterior spines, rostrate long-shafted uncini and spinose or smooth capillaries; seg-
ments usually elongated (bamboo-worms) MALDANIDAE
69b (68a). Setae include bilimbate and spatulate kinds, long-handled uncini and companion-setae; segments
not prolonged SABELLONGIDAE
70a (68b). Body separated into two regions with different kinds of setae in a thoracic and abdominal region
( NOTE: Regions may sometimes also be definable on parapodial features) 71
70b (68b). Body not separated into regions; setal distribution and parapodial shapes grade along the body . . 73
71a (70a). Thorax with lateral parapodia, abdomen with both noto- and neuropodia in dorsal positions
. . . . . . . . . . . . . .......................................................... ORBINIIDAE
71b (70a). Parapodia lateral in all parts of the body; notopodia often reduced in posterior segments and neuro-
podia may form nearly complete cinctures 72
72a (71b). Slender capillary setae in thorax and sometimes in the first few abdominal segments only; branchiae,
if present, retractable filaments on the abdomen CAPITELLIDAE
72b (71b). Slender capillary setae present on anterior, median and sometimes posterior parts of the body, in-
cluding the branchial region; branchiae non-retractable, bushy or simple filaments ARENICOLIDAE
73a (70b). Anterior end with complex jaw-apparatus 74
73b (70b). Jaw-apparatus absent 76
74a (73a). Each jaw consisting of a series of small denticles DORVILLEIDAE
74b (73a). Each jaw consisting of a single piece 75
75a (74b). Hooded hooks present in at least some setigers; one pair of maxillary carriers .. LUMBRINERIDAE
75b (74b). Hooded hooks absent; three maxillary carriers ARABELLIDAE
76a (73b). Branchiae present on maximally 15-20 segments, starting from one of setigers 4-10
. . .............................................................. PARAONIDAE (part)
76b (73b). Branchiae, if present, either limited to the extreme anterior end, or found scattered over a large part
of the body 77
77a (76b). Setae include anterior spines, rostrate long-shafted uncini and spinose or smooth capillaries; segments
usually elongated (bamboo-worms) MALDANIDAE
77b (76b). Setal distribution otherwise; segments rarely elongated 78
78a (77b). Setae include simple capillaries and simple bifid falcigers in both noto- and neuropodia in a long
region of the body QUESTIDAE
78b (77b). Simple bifid falcigers absent 79
79a (78b). Prostomium an elongated cone, usually more than twice as long as wide, nearly always articulated 80

79b (78b). Prostomium less than twice as long as wide, may be bluntly conical, rounded or truncate, never
articulated 81
80a (79a). Eversible pharynx with four jaws; parapodia either all uniramous or all biramous
. . . . . . . . ......................................................... GLYCERIDAE (part)
80b (79a). Eversible pharynx with more than four jaws; anterior parapodia uniramous, posterior ones bi-
ramous GONIADIDAE
8la (79b). Parapodia rounded lobes with large, easily dehiscent cirri; pelagic TYPHLOSCOLECIDAE
81b (79b). Parapodia low folds or blunt, button-shaped projections; cirri, if present, cirriform 82
82a (81 b). Composite falcigers present in anterior setigers ACROCIRRIDAE (part)
82b (81b). All setae simple 83
83a (82b). Neurosetae uncinal in structure BOGUEIDAE
83b (82b). Unciniabsent 84
84a (83b). All setae simple capillaries; branchiae cirriform, pectinate or absent; prostomium entire, pointed
or rounded OPHELIIDAE
84b (83b). Furcate and acicular setae usually present; branchiae, if present, limited to the anterior end and strongly
arborescent; prostomium T-shaped or bifid SCALIBREGMIDAE
85a (58b). Prostomium with a median antenna 86
85b (58b). Prostomium without appendages 88
86a (85a). Branchiae present on maximally 15-20 segments first starting from one of setigers 4-10
. . . . . ........................................................... PARAONIDAE (part)
86b (85a). Branchial distribution otherwise or branchiae absent 87
87a (86b). Notopodial cirri flask-shaped in some setigers, plumose setae present POECILOCHAETIDAE
87b (86b). Notopodial cirri cirriform or foliose, plumose setae absent SPIONIDAE (part)
88a (85b). With two pairs of tentacular cirri; setae composite 89
88b (85b). Without tentacular cirri; setae simple 90
89a (88a). Setal appendages long and slender; pelagic IOSPILIDAE
89b (88a). Setal appendages short; parasitic CALAMYZIDAE
90a (88b). With multiple series of small, long-shafted uncini in the neuropodia OWENIIDAE (part)
90b (88b). Uncini in single rows or absent 91
91 a (90b). Prostomium flattened and spatulate; as wide as the widest part of the body MAGELONIDAE
91b (90b). Prostomium not flattened; distinctly narrower than the widest part of the body 92
92a (91 b). Parapodia inconspicuous; abdominal segments prolonged with setae forming complete cinctures around
the body HETEROSPIONIDAE
92b (91 b). Parapodia well developed; abdominal segments usually not prolonged; setae limited to lateral tufts 93
93a (92b). All parapodia biramous, except possibly the first one 94
93b (92b). Median parapodia uniramous TROCHOCHAETIDAE
94a (93b). Uncini absent; notopodia with acicula, but without setae; true branchiae absent APISTOBRANCHIDAE
94b (93b). Uncini normally present, at least in the neuropodia; notopodia with setae in addition to acicula; branchiae
present or absent SPIONIDAE (part)
ORDER ORBINIIDA
Prostomium without appendages; maximally two Prostomium without appendages, one or two asetigerous
asetigerous anterior segments present; no additional segments present anteriorly; no peristomial appendages.
cephalized segments present. Palps absent; eversible A saclike proboscis present. All setae simple, including
pharynx either an axial sac or a ventrolateral pad. capillaries, simple hooks and sometimes brush-topped,
Parapodia biramous; all setae simple, including cap- bifid or furcate setae.
illary setae and usually acicular spines and serrated or The orbiniids have been the subject of several com-
spinose setae. prehensive studies, including Hartman (1957), Petti-
bone (1957a) and Day (1973). The major generic
FAMILY ORBINIIDAE HARTMAN 1942 groupings appear clear and were reviewed by Day
Orbiniida with lateral parapodia in a thoracic region (1973) for the subfamily Orbiniinae. The subfamily
and, usually, dorsal parapodia in an abdominal region. Protoariciinae has yet to be reviewed in detail.

E
FIGURE2. (A), Family ORBINIIDAE, Orhinia johnsoni, anterior end, after Hartman 1969, 5x; (B), transverse section of the
abdomen of the above, 18x; (c), Family PARAONIDAE, Cirrophores sp., anterior end, diagrammatic, about 22x; (D), Family
QUESTIDAE, Questa caudicirra, posterior seta, after Hartman 1966, 54x; (E) anterior end of the above, 24x.
KEY TO GENERA
Ia. Two asetigerous anterior segments 2

lb. Asingleatrou sgmen2a(I).
Branchiae absent Orbiniella
2b (la). Branchiae present 3
3a (2b). Branchiae present on all but a few anterior segments Protoariciella
3b (2b). Branchiae limited to abdominal segments 4
4a (3b). Only crenulated capillaries present Scoloplella
4b (3b). Crenulated capillaries and other kinds of setae present 5
5a (4b). Abdominal neurosetae include swan-shaped hooks Proscoloplos
5b (4b). Swan-shaped hooks absent 6
6a (5b). Abdominal neurosetae all crenulated capillaries; prostomium rounded Protoaricia
6b (5b). Abdominal neurosetae include acicular hooks; prostomium pointed Schroederella
7a(lb). Branchiae absent Microrbinia
7b (lb). Branchiae present on at least some segments 8
8a (7b). Prostomium rounded or truncate Naineris
8b (7b). Prostomium more or less pointed 9
9a (8b). All thoracic parapodia with only slender, pointed setae 10
9b (8b). Some thoracic neuropodia with setae of another kind 1l
l0a (9a). Abdominal neuropodia with acicular spines Berkeleya

lob (9a). Acicular spines absent in abdominal neuropodia Haploscoloplos

I la (l0b). Thoracic neuropodia of two abruptly different kinds 12
I lb (l0b). Thoracic neuropodia not abruptly different 13
12a (1l a). Anterior three thoracic neuropodia with bristle-tipped setae Califia
12b (11 a). Posterior thoracic neuropodia with thick, modified spines associated with a glandular pouch .. Phylo
13a (I 1b). Some thoracic segments with rows of papillae along the ventrum, papillae sometimes also on parapodial
postsetal ridges Orbinia
13b (I lb). Without rows of papillae on the ventrum 14
14a (13b). Median and posterior abdominal neuropodia with thick acicula, projecting from the parapodial lobes;
branchiae usually present from fifth or sixth segment Scoloplos (Leodamas)
l4b (13b). Thick projecting acicula absent; branchiae usually not present before tenth segment
. . . . . . . . . . . . . . . ..........................................................(Scoloplos)
Generic Definitions include crenulated capillaries, hooks and intermediate

forms. Abdominal notosetae crenulated capillaries and
Berkeleya Hartman 1971, B. profunda Hartman 1971; sometimes furcate setae.
only species.
ORBINIINAE with pointed prostomium, abdominal Orbinia Quatrefages 1865, Aricia cuvierii Audouin and
branchiae and all thoracic setae crenulated capillaries. Milne Edwards 1833c; 16 species.
Twelve thoracic segments; transition from thorax to ORBINIINAE with pointed prostomium and first
abdomen abrupt. Abdominal notopodia with crenulated pair of branchiae on setiger 5-9. Posterior thoracic
capillaries and furcate setae; abdominal neuropodia parapodia with several accessory papillae and numerous
with crenulated capillaries and acicular spines. ventral papillae; with a combined total of at least five
papillae on each segment. Thoracic neurosetae include
Califia Hartman 1957, C. candida Hartman 1957; 4
hooks (or subuluncini) and crenulated capillaries.
species.
Furcate and capillary setae present in abdominal
ORBINIINAE with pointed prostomium, branchiae
notopodia.
not present before segment 8; posterior thoracic neuro-
podia with two or fewer papillae; ventral papillae ab- Orbiniella Day 1954, O. minuta Day 1954; 4 species.
sent. Thoracic neurosetae include anterior brush-topped PROTOARICIINAE with rounded or pointed pros-
setae, crenulated capillaries and in some cases blunt tomium and without branchiae. Thoracic setae all
hooks. camerated or crenulated capillaries; abdominal setae
include crenulated capillaries, acicular setae and some-
Haploscoloplos Monro 1933, Scoloplos cylindrifer
times furcate setae.
Ehlers 1905; 13 species.
ORBINIINAE with pointed prostomium, branchiae Phylo Kinberg 1866b, P. felix Kinberg 1866b; 19
not present before setiger 9; posterior thoracic neuro- species.
podia with two papillae and maximally two ventral ORBINIINAE with pointed prostomium and branchiae
papillae present; never more than four papillae com- first present from setiger 5-7. Posterior thoracic para-
bined. All neurosetae crenulated capillaries; notosetae podia with several accessory papillae; numerous ventral
crenulated capillaries and in some cases furcate setae. papillae present. At least five papillae in combined
total present on a segment. Thoracic neurosetae include
Microrbinia Hartman 1965, M. linea Hartman 1965; crenulated capillaries, heavy hooks and heavy spear-
only species.
shaped setae.
ORBINIINAE with conical prostomium; branchiae
and accessory papillae absent. Separation between Proscoloplos Day 1954, P. cygnochaetus Day 1954;
thorax and abdomen indistinct. First notosetae smooth 2 species.
capillaries; other notosetae camerated capillaries and PROTOARICIINAE with rounded prostomium and
posterior notosetae curved and serrated acicular spines. stout branchiae present from setiger 8. Transition from
Neurosetae similar. thorax to abdomen indistinct. Setae include crenulated
capillaries and one or two swan-shaped hooks in pos-
Naineris Blainville 1828, Nais quadricuspida Fabricius terior neuropodia.
1780; 18 species.
ORBINIINAE with rounded or truncate prostomium, Protoaricia Czerniavsky 1881a, Aricia oerstedii
branchiae first present from any segment from 2-23. Claparede 1864; 2 species.
Maximally two accessory papillae on posterior para- PROTOARICIINAE with rounded prostomium and
podia, ventral papillae absent. Thoracic neurosetae branchiae limited to abdominal segments. Thoracic

neurosetae include crenulated capillaries, hooks and Invalid Genera

subuluncini; abdominal neurosetae all crenulated
capillaries. Alcandra Kinberg 1866b, see Scoloplos (Leodamas)
Anthostoma Schmarda 1861, see Naineris
Protoariciella Hartmann-Schroder 1962, P. uncinata Aricia Savigny 1820, see Orbinia
Hartmann-Schroder 1962; 3 species. Branchetus Chamberlin 1919c, see Scoloplos (Leo-
PROTOARICIINAE with pointed prostomium and damas)
branchiae present on all but a few anterior segments Clytie Grube 1855, indeterminable
( from setiger 3 in type species). Thoracic notosetae Gisela MUller 1858, indeterminable
all are crenulated capillaries; acicular setae present Labotas Kinberg 1866b, see Scoloplos
in posterior notopodia. Neurosetae include thick hooks Lacydes Kinberg 1866b, see Naineris
with three teeth; slender acicular setae with flat teeth, Naidonereis Malmgren 1867, error for Naineris
crenulated capillaries and thick smooth spines in pos- Theodisca Muller 1858, see Naineris
terior segments. Theostoma Eisig 1914, see Protoaricia
Venadis Castelnau 1842, see Orbinia
Schroederella Laubier 1962, S. pauliana Laubier 1962;
2 species.
PROTOARCIINAE with strongly pointed pros- Incertae Sedis
tomium and branchiae present on the abdomen. Noto-
setae include crenulated capillaries and acicular spines Falklandiella Hartman 1967, F. annulata Hartman
in far posterior segments. Neurosetae include crenu- 1967; only species.
lated capillaries and acicular hooks. Resembles the orbiniids in setal structures, but ap-
pears to differ in other features. Prostomium a simple
Scoloplella Day 1963, S. capensis Day 1963; only
triangular lobe; two asetigerous segments present. Para-
species.
podia biramous with very strongly reduced parapodial
PROTOARICIINAE with pointed prostomium and lobes. Notosetae all slender and camerated; most neuro-
branchiae present on the abdomen. All setae crenulated setae similar, but shorter; in addition heavy acicular
capillaries. spines present in posterior neuropodia.
Scoloplos Blainville 1828, Lumbricus armiger O.F.
Muller 1776; 38 species.
ORBINIINAE with pointed prostomium and branchiae
FAMILY PARAONIDAE CERRUTI 1 909
first present from setiger 5 or later. Accessory papillae Body long and slender with lateral parapodia. Pros-
never exceeding four, including both ventral and para- tomium with a single antenna or antennae absent.
podial kinds. Thoracic neurosetae include blunt hooks Branchiae present on a limited number of median
and crenulated capillaries (may be absent). Abdominal setigers in most species. All setae simple, including
notosetae include crenulated capillaries, furcate setae capillaries and various, usually postbranchial, hooks
and spines. or otherwise modified setae.
Subgenus Scoloplos: Species with first branchiae The key given below is based on the important re-
on setigers 8-10 or later; without emerging notacicula vision by Strelzov (1973); some features are taken
in posterior setigers. from Fauchald (1972). Strelzov recognized only two
Subgenus Leodamas: Species with first branchiae on genera with prostomial antennae, Cirrophorus and
setiger 5-6; with heavy emerging notacicula in pos- Aricidea; the latter with a series of subgenera; these
terior setigers. subgenera are here considered distinct genera.
Key to Genera
Ia. Setae present from the first segment Aparaonis
lb. Setae present from the second segment 2
2a (1 b). Modified setae notopodial Cirrophorus
2b (Ib). Modified setae, if present, neuropodial 3
3a (2b). Prostomium with a median antenna 4
3b (2b). Prostomium without a median antenna 7
4a (3a). Modified setae absent Aedicira
4b (3a). Modified setae present 5
5a (4b). Modified setae either pseudocomposite or curved with a subterminal arista on concave side of the
shaft Aricidea

5b (4b). Modified setae otherwise 6

6a (5b). Modified setae not greatly different from the capillary setae, but somewhat thicker and more abruptly
tapering and with a long arista Allia
6b (5b). Modified setae distally strongly curved, or bifid, either with numbers of long aristae or bearded .
Acesta
7a (2b). Nuchal organ on posterior part of prostomium, prostomium with several transverse bands of
cilia Paraonis
7b (2b). Nuchal organ on the reduced peristomium; with a single preoral ciliary band or prostomial ciliary
bands absent 8
8a (7b). Modified setae absent Paraonella
8b (7b). Modified setae present 9
9a (8b). Three prebranchial setigers; terminal prostomial sense organs absent Sabidius
9b (7b). More than three prebranchial setigers; terminal prostomial sense organs present Tauberia
Generic Definitions Nuchal organs prostomial; median antenna absent.

Modified setae neuropodial.
Acesta Strelzov 1973, Aricidea catherinae Laubier
1967a; 10 species. Sabidius Strelzov 1973, Paraonis cornatus Hartman
Median antenna present. Modified setae neuropodial; 1965; only species.
each strongly curved distally, or bifid with several Nuchal organs peristomial; median antenna absent.
aristae or with a short stiff beard. Modified setae neuropodial. Prostomial terminal sense
organs absent. Three prebranchial setigers.
Aedicira Hartman 1957, Aricidea Pacifica Hartman
1 944c; only species. Tauberia Strelzov 1973, Aonides gracilis Tauber 1879;
Median antenna present. All setae slender capillaries; 10 species.
those in the neuropodial fascicles often shorter than Nuchal organs peristomial; median antenna absent.
the notopodial ones. Modified setae neuropodial. Prostomial terminal sense
organs present. More than three prebranchial setigers
Allia Strelzov 1973, Aricidea albatrossae Pettibone
present.
1957b; 13 species.
Median antenna present. Modified setae slightly
thicker than the non-modified capillary setae; somewhat Taxonomic Notes
more abruptly tapering; or tapering abruptly near the The generic subdivision given above, differs mark-
middle, giving the outer part of the seta the appearance edly from the more traditional pattern summarized
of a long, smooth arista. by Fauchald (1972). Strelzov (1973) introduced sev-
Aparaonis Hartman 1965, A. abyssalis Hartman 1965; eral new taxonomic characters to the ones previously
only species. in common usage. The presence of prostomial ciliary
Setae present in the first segment; three pairs of bands, of prostomial terminal sense organs and the local-
branchiae on segments 2-4. All setae simple capil- ization of the nuchal organs must now be investigated.
laries. Similar characters have been used in other families,
and it is anticipated that they will gain acceptance also
Aricidea Webster 1879b, A. fragilis Webster 1879b; in the treatment of the paraonids.
7 species.
Median antenna present; modified setae either pseudo- Invalid Genus
composite or recurved with a subterminal arista arising
from the concave side of the shaft. Levinsenia Mesnil 1897, see Paraonis.
Several other genera, such as Paraonides, Paradoneis
Cirrophorus Ehlers 1908, C. branchiatus Ehlers 1908; and others used in the traditional sub-division of the
10 species. family, may be invalid. Until the value of Strelzov's
Median antenna present or absent; modified setae system has been tested, I hesitate to treat these genera
present in the postbranchial notopodial fascicles. as invalid.
Paraonella Strelzov 1973, Paraonides nordica Strelzov
1968a; 4 species. FAMILY QUESTIDAE HARMAN 1966a
Nuchal organs on the peristomium; median antenna
Body long and slender. Prostomium without append-
absent. Modified setae absent.
ages. One asetigerous anterior segment present. Phar-
Paraonis Cerruti 1909, Aonides fulgens Levinsen 1884; ynx with a muscular pad. Parapodia biramous with
2 species. reduced lobes. Branchiae and anal cirri present or

absent. Setae include serrated capillaries, bidentate from the polychaetes (Fauchald 1974a). However, the
hooks and trifurcate spines. questids have several different kinds of setae; a feature
As remarked by Hobson (1970) the questids have the rarely found among the oligochaetes. It is quite pos-
gonads limited to a few segments; this generally is sible that the family should be considered among the
considered the key feature separating the oligochaetes oligochaetes.
Key to Genera
Ia. Setae include capillaries and bidentate hooks; posterior cirriform branchiae present Questa
lb. Setae include capillaries, bidentate hooks and trifurcate spines; branchiae absent Novaquesta
Generic Definitions (1971); it now is divided into two sub-families with

three genera in all; each with only a few species.
Novaquesta Hobson 1970, N. trifurcata Hobson 1970;
only species.
Questids with serrated capillary setae, bidentate
hooks and trifurcate spines with the middle tine shorter
and slenderer than the other; branchiae and anal cirri
absent.
Questa Harman 1966a, Q. caudicirra Hartman 1966a;

only species.
Questids with serrated capillary setae and bidentate
hooks. Branchiae present on posterior segments, anal
cirri present.
ORDER CTENODRILIDA
Prostomium without appendages; palps absent. At
l east one asetigerous anterior segment present. Pro-
boscis a ventral muscular pad. Parapodia uni- or bi-
ramous; all setae simple. Parapodial lobes absent.
Members of this order are very small, generally
grub-shaped polychaetes that tend to turn up in mass-
culture in aquaria more frequently than in the field
(especially true for the ctenodrilids). Specimens of
Ctenodrilus also have turned up associated with Fla-
belliderma commensalis at Santa Catalina Island and
have the same dark purple pigmentation on sea urchins
as these commensals do.
FAMILY CTENODRILIDAE KENNEL 1882
Body short or long, prostomium without appendages,

palps absent. Pharynx a ventral muscular pad. Some
anterior segments may be fused and asetigerous; some
segments with median dorsal cirri; paired lateral
A
branchiae present in some juveniles. Setae in double
fascicles; parapodial lobes absent.
FIGURE 3. (A), Family CTENODRILIDAE, Crenodrilus ser-
ratus, modified after Hartman 1 944, 284x; (B), Family PARER-
This small family of tiny polychaetes recently has GODRILIDAE, Stygocapitella subterranea, after Hattmann-
been reorganized and redefined by Hartmann-Schroder Schroder 1971, about 300x.
Key to Genera
Ia. Body short and grublike, with few segments, unpaired dorsal cirri and branchiae absent
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . CTENODRILINAE Ctenodrilus
lb. Body long and slender, with at least 15 segments; unpaired dorsal cirri on one or two anterior seg-
ments or larval branchiae present RHAPHIDRILINAE 2

2a (lb). Unpaired dorsal cirri present; branchiae absent Zeppelinia

2b (Ib). Unpaired dorsal cirri absent; branchiae present Raphidrilus
Generic Definitions Setae include capillaries and short, thicker dentate or

smooth spines. One or two unpaved dorsal cirri pres-
Ctenodrilus Claparede 1863, Parthenope cirrata Schmidt ent anteriorly.
1857; 2 species.
CTENODRILINAE. Body with maximally 15 seg- Invalid Genera
ments; one achaetous segment present. Setae thick, Monostylos Vejdovsky 1884, see Zeppelinia
marginally coarsely dentate or smooth. Parthenope Schmidt 1857, see Ctenodrilus
Rhaphidrilus Monticelli 1910, R. nemasoma Monticelli
1910; only species. FAMILY PARERGODRILIDAE REISINGER 1960
RHAPHIDRILINAE. Body long and slender, with Small, grub-shaped worms with at least one anterior
at least 18 segments; one asetigerous segment pres- asetigerous segment. Prostomium without appendages;
ent. Setae include capillaries and genital spines in parapodial lobes absent; setae present in single fascicles,
the adult males. Paired branchiae present in the juve- each with one or a few setae only. Setae all simple.
nile stages. The parergodrilids are very small worms, living either
Zeppelinia Vaillant 1890, Ctenodrilus monostylos Zep- interstitially in shallow marine sands (Stygocapitella)
pelin 1883; 5 species. or in rotting terrestrial plant material (Parergodrilus).
RHAPHIDRILINAE. Body long and slender, with The family is poorly known and understood; the revi-
at least 18 segments; asetigerous segments absent. sion by Hartmann-Schrdder (1971) is followed here.
Key to Genera
la. Body not translucent, with 10-11 setigers; setae limbate and furcate Stygocapitella
lb. Body translucent, 8-9 setigers present; all setae smooth rods Parergodrilus
Generic Definitions
Parergodrilus Reisinger 1925, P. heideri Reisinger

1925; only species.
One anterior and no posterior asetigerous segments;
eight or nine setigers present. Setae all smooth rods,
except copulatory hooks in males.
Stygocapitella Knollner 1934, S. subterranea Knollner

1934; only species.
Two anterior and two posterior asetigerous segments,
ten or 11 setigers present. Setae include smooth, short
bilimbate setae, smooth bilimbate setae with long,
whiplike tips and short furcate setae with two accessory
teeth in the crotch.
ORDER PSAMMODRILIDA
Prostomium without appendages, palps absent. At

least one asetigerous anterior segment present. Noto-
podia in a median region strongly prolonged and sup-
ported by acicula. Parapodia biramous; neuropodial
tori present in one form, otherwise parapodial lobes
absent (except for the cirri mentioned above); all setae
simple.
A
FAMILY PSAMMODRILIDAE SWEDMARK 1952
FIGUae 4. (A), Family PSAMMODRILIDAE, Psammodril-
Small, grub-shaped worms. Pro- and peristomium oides fauveli after Swedmark 1958, about 50x; (B), Family
without appendages; palps absent. Parapodia, except COSSURIDAE, Cossura brunnea, after Fauchald 1972, 12.5x.

the thoracal dorsal cirri reduced; abdominal neuro- The first form described, Psammodrilus balanoglos-
podial tori present in one form. soides Swedmark, is unique in that the eversible pharynx
As remarked by Swedmark (1958), these small inter- is formed from the longitudinal body muscles.
stitial worms are very isolated among the polychaetes.
Key to Genera
Ia. Dorsal cirri decreasing evenly in length from the first to the sixth thoracic segment; abdominal
uncinigers with several uncini Psammodrilus
lb. Dorsal cirri increasing in length from first to fourth segment; those of segments five and six very
short; single uncini present in abdominal neuropodia Psammodriloides
Generic Definitions rata Webster and Benedict (1887) with a total of about
fifteen species. Two of these differ rather sharply
Psammodriloides Swedmark 1958, P. fauveli Swed-
from the others and may deserve separate generic
mark 1958; only species.
standing.
Body with two distinct regions: head and trunk;
six thoracic setigers, each with a dorsal cirrus sup-
ported by an aciculum. Ten abdominal setigers; each ORDER SPIONIDA
with a single uncinus in each neuropodium. Eversible
pharynx absent. Prostomium distinct, without appendages or with an
occipital antenna. A pair or two groups of grooved
Psammodrilus Swedmark 1952, P. balanoglossoides feeding palps usually present anteriorly, either on the
Swedmark 1952; only species. prostomium or on an anterior segment. Paired tentacular
Body in three regions: head, pharyngeal region and cirri sometimes present at the base of the palps. Pharynx
trunk. Six thoracic setigers with dorsal cirri supported either axial or as a ventral pad. Parapodia well devel-
by acicula; up to about thirty abdominal setigers; each oped or reduced. All setae simple.
with several uncini in each neuropodium. Pharyngeal
apparatus present.
ORDER COSSURIDA
Prostomium without appendages; a single peristomial

asetigerous segment present; a single median palp
present on the dorsum of an anterior setiger (usually
between setigers 3 and 6). Proboscis a ventral pad.
Parapodia biramous, with very low parapodial lobes;
all setae simple.
FAMILY COSSURIDAE DAY 1 963
Prostomium without appendages, a single median

palp present dorsally on one anterior setiger; proboscis
a ventral pad. Parapodia biramous, with reduced para-
podial lobes. All setae simple, including bilimbate or
hirsute setae in two or more fascicles; thick spines
and capillary setae present in the abdomen of some
forms.
Cossurids are common in sand and especially in
deep slope and abyssal muds. They are burrowers,
and apparently feed on detritus with the help of the
pharynx. The palp in this case appears to be sensory, FIGURE 5. (A), Family APISTOBRANCHIDAE, Apistobran-
and perhaps may additionally be respiratory in func- chus tullbergi, ventral view, after Pettibone 1963, lox; (B),
tion since it is well-equipped with blood-vessels dorsal view of the above, lox; (C), Family TROCHOCHAE-
TIDAE, Trochochaetus multisetosum, lateral view, after
(Fauchald in preparation). Pettibone 1 963, 1 Ox; (D), dorsal view of the above, I Ox; (E),
The only currently described genus is Cossura Web- Family SPIONIDAE, Boccardia proboscidea, modified after
ster and Benedict, (1887) with genotype C. longocir- Hartman 1 969, 25x.

Key to Suborders
Ia. Body separated into at least two distinct regions; uncini present Chaetopteriformia
lb. Body rarely separated into distinct regions, uncini always absent 2
2a (I b). Palps at the junction of the pro- and peristomium; parapodial lobes well developed at least in some
setigers Spioniformia
2b (lb). Palps post-peristomial in origin; parapodial lobes poorly developed Cirratuliformia
Suborder Spioniformia FAMILY SPIONIDAE GRUBE 1850
Palps on the peristomium or at the junction between Spioniforms with body elongated. Prostomium an-
the pro- and peristomium; eversible pharynx either a teriorly blunt, with frontal horns, or pointed; an oc-
ventral pad or an axial sac. cipital papilla may be present, other appendages absent.
Palps at the postectal comers of the prostomium. Para-
FAMILY APISTOBRANCHIDAE
podia biramous, parapodial lobes cirriform or foliose,
MESNIL AND CAULLERY 1898
never serrated. All setae simple, including capillaries
Spioniforms with the anterior end inflated and the and bi- or multidentate, hooded or non-hooded hooks.
rest of the body cylindrical. Prostomium without ap- Spionids are very common in all environments.
pendages, but with palps attached at the postectal Some forms are burrowers (Polydora, Boccardia et
margins. Parapodia biramous, parapodial lobes mostly al.) in calcareous substrata or in rock; others build
cirriform, but serrated postsetal lobes present in some permanent tubes in soft substrata and some forms com-
setigers. All setae simple, mainly capillaries. bine the two activities. Numerous forms are free-
The apistobranchids are known for one genus, living in sands and muds. Species of some genera,
Apistobranchus Levinsen 1883, with type species most frequently perhaps of Prionospio SENSU LATU and
Aricia tullbergi Theel 1879. A total of three species of the Polydora/Boccardia complex, very likely are
currently are recognized. present in any kind of benthic marine soft-bottom
Apistobranchids are in fact not as rare as the low sample taken anywhere on the globe. Members of the
number of species should indicate; they are, as some genus Aonides are dominant forms in sandy beaches
of their spionid relatives, not limited to a tubicolous in temperate and warm areas of the world (Foster 1971).
existence, but will feed from loosely constructed bur- The spionids usually lose their palps, and frequently
rows. Most commonly found in shelly sands, sands most of their branchiae in fixation. This is especially
and muds. true in those genera where the structure of the branchiae
is important for species separation. Attempts at identifi-
Invalid Genera cation of these taxa on incomplete material should be
Ethocles Webster and Benedict 1887, see Apisto- avoided; the family is speciose and serious errors are
bronchus easily introduced into the literature, if care is not taken
Skardaria Wesenberg-Lund 1951, see Apistobranchus at all identificatory levels.
Key to Genera
Ia. Each branchia with several (maximally seven) processes in median parapodia Polybranchia
2
1b. Branchia, if present, single process
2a (I b). One setiger with strongly modified setae 3
2b (lb). Setae change gradually along the body, no segment with remarkably different setae (except large
hooks in the first setiger) 8
3a (2a). Setiger 16 modified Morants
4
3b (2a). Either setiger 4 or 5 modified
4a (3b). Setiger 4 modified Polydorella
4b (3b). Setiger 5 modified 5
5a (4b). Neuropodial uncini distally trifid Tripolydora
5b (4b). Neuropodial uncini distally bifid or entire 6
6a (5b). Branchiae first present from setiger 2 Boccardia
6b (5b). Branchiae first present posterior to the modified segment 7
7a (6b). Setae in modified segment arranged in a horseshoe-shaped series Pseudopolydora
7b (6b). Setae in the modified segment in a straight line or a small patch Polydora
8a (2b). Branchiae absent; first neuropodia with stout hooks Spiophanes

8b (2b). At least one pair of branchiae present; first neuropodia without stout hooks 9
9a (8b). Branchiae present on nearly the whole body 10
9b (8b). Branchiae limited to less than half the length of the body 19
l0a (9a). Branchiae present from setiger 1 Il
l 0b (9a). Branchiae present from setiger 2 14
I 1a (l0a). Accessory branchiae on some segments Dispio
I I b (l0a). Accessory branchiae absent 12
12a (I lb). Posterior notopodia with uncini Marenzelleria
12b (I lb). Notosetae all capillaries 13
13a (12b). Prostomium anteriorly rounded Spio
13b (12b). Prostomium with laterofrontal horns Malacoceros
14a (lob). Posterior notopodia with uncini Scolelepis
14b (lob). All notosetae capillaries 15
15a (14b). Prostomium with laterofrontal horns 16
15b (14b). Prostomium anteriorly blunt or pointed 17
1 6a (15a). Branchiae partially fused to the notopodial postsetal lobes Rhynchospio
16b (15a). Branchiae completely separated from the notopodial postsetal lobes Mesospio
1 7a (15b). Occipital antenna absent Microspio
1 7b (15b). Occipital antenna present 18
18a (17b). Branchiae partially fused to the notopodial postsetal lobes Pseudomalacoceros
18b (17b). Branchiae completely separated from the notopodial postsetal lobes Laonice
19a (9b). A single pair of branchiae present Streblospio
1 9b (9b). At least two pairs of branchiae present 20
20a (19b). Branchiae concentrated in a medio-posterior region, except in the males, where an additional single
pair is present on the second setiger Pygospio
20b (19b). Branchiae concentrated near the anterior end only 21
21 a (20b). Peristomium forms large lateral wings on either side of the prostomium 22
21b (20b). Peristomium does not form lateral wings on the sides of the prostomium 24
22a (21 a). Three pairs of pinnate branchiae present from setiger I Paraprionospio
22b (21 a). Branchiae otherwise, first present from setiger 2 23
23a (22b). Two to four pairs of branchiae present Aquilaspio
23b (22b). At least six pairs of branchiae present Minuspio
24a (21b). Prostomium anteriorly pointed Aonides
24b (21b). Prostomium anteriorly blunt or with frontal horns 25
25a (24b). Prostomium with frontal horns Scolecolepides
25b (24b). Prostomium anteriorly rounded 26
26a (25b). Branchiae two pairs on setigers 3-4 Anaspio
26b (25b). Four pairs of branchiae from setiger 2 27
27a (26b). Three first pairs of branchiae cirriform, the last pinnate Apoprionospio
27b (26b). Branchiae cirriform or pinnate in another pattern Prionospio
Generic Definitions free from the notopodial postsetal lobes. Anterior se-
tigers with capillary setae only; posterior parapodia
Anaspio Chamberlin 1920, A. boreus, Chamberlin with uncini in both rami. Neuropodial postsetal lobes
1920; only species. smooth.
Prostomium anteriorly rounded. Two pairs of bran- Apoprionospio Foster 1969, A. dayi Foster 1969; 5
chiae on setigers 3 and 4; free from the notopodial species.
postsetal lobes. Notosetae all capillaries (in 36 setigers); Prostomium anteriorly blunt; peristomium without
neuropodia with uncini from setigers 10-11. Anterior lateral wings. Four pairs of branchiae from setiger 2;
parapodial lobes very large and foliose. first three cirriform, fourth pinnate. All anterior setae
Aonides Claparede 1864, Nerine oxycephala Sars 1862; capillaries; posteriorly uncini in both rami. Neuropodial
postsetal lobes large and foliose in setiger 2.
7 species.
Prostomium anteriorly pointed. Branchiae from Aquilaspio Foster 1971, Prionospio sexoculata Augener
setiger 2, present on anterior end only; completely 1918; 2 species.

Prostomium anteriorly blunt; peristomium with large Minuspio Foster 1971, Prionospio cirrifera Wiri;n 1883;
lateral wings; first setiger more or less reduced. Two 7 species.
to four pairs of branchiae from setiger 2. All anterior Prostomium anteriorly blunt; peristomium with large
setae capillaries; posterior setigers with uncini in lateral wings. Branchiae present from setiger 2 to about
both rami. setiger 40, all cirriform. Anterior setae all capillaries in
both rami; posterior end with uncini in both rami.
Boccardia Carazzi 1895, Polydora (Leucodore) poly-
branchia Haswell 1885; 15 species. Morants Chamberlin 1919a, M. duplex Chamberlin
Prostomium anteriorly blunt or bifid. Branchiae 1919a; only species.
present from setiger 2. Fifth setiger modified with First four setigers with branchiae; fused to the noto-
strong setae; in other setigers, all notopodia with podial postsetal lobes; setiger 16 with strongly modified
capillaries only; neuropodia posteriorly with uncini. setae. Curved hooks present in the first notopodia; all
other notosetae capillary. Anterior neurosetae capillary,
Dispio Hartman 1951a, D. uncinata Hartman 1951a; posterior ones uncini.
5 species.
Prostomium anteriorly blunt; peristomium enfolding Paraprionospio Caullery 1914b, Prionospio pinnata
sides of prostomium. Branchiae from setiger 1 to the Ehlers 1901; 5 species.
end of the body; at least partially fused to the noto- Prostomium anteriorly rounded; peristomium with
podial postsetal lobes; accessory branchiae present in l arge lateral wings. Second segment asetigerous. Three
some setigers. Notosetae all capillaries; neurosetae pairs of pinnate branchiae present from setiger 1; para-
include capillaries and uncini in median and posterior podia of first setiger well developed. All anterior setae
setigers. capillaries; posterior parapodia with uncini in both rami.
Laonice Malmgren 1867, Nerine cirrata Sars 1850; Polybranchia Potts 1928, P. foxi Potts 1928; only
15 species. species.
Prostomium anteriorly rounded; occipital antenna Prostomium with frontal horns. Branchia present
present. Branchiae present from setiger 2 to the middle from setiger 1; each branchia with maximally six to
of the body; completely free from the notopodial seven processes in median setigers. Capillary setae
postsetal lobes. Notosetae all capillaries; neurosetae absent; all setae uncini in both rami.
include capillaries and uncini. Genital pouches in at
Polydora Bosc 1802, P. cornuta Bose 1802; 65 species.
least some setigers.
Prostomium anteriorly blunt or bifid. Branchiae not
Malacoceros Quatrefages 1843, Spio vulgaris Johnston present before setiger 6. Setiger 5 with strongly modified,
1827; 10 species. stout setae. Other setae include notopodial capillaries,
Prostomium with lateral horns. Branchiae present sometimes also with simple posterior spines. Neuropodial
from setiger 1, partially fused to the notopodial post- uncini present from setiger 7-10 in most species.
setal lobes. Notosetae all capillaries; neurosetae include
Polydorella Augener 1914, P. prolifera Augener 1914;
capillaries and uncini in posterior setigers.
2 species.
Marenzelleria Mesnil 1896, M. wireni Augener 1913b; Prostomium anteriorly blunt. Branchiae from setiger 6.
only species. Fourth setiger modified with large, stout setae. Notosetae
Prostomium anteriorly rounded. Branchiae present all capillaries, except in setiger 4; neuropodia with capil-
from setiger 1, partially fused to the notopodial post- laries, but with uncini present from setiger 6.
setal lobes. Anterior setigers with capillary setae in
Prionospio Malmgren 1867, P. steenstrupi Malmgren
both rami; posteriorly uncini in both rami.
1867; 36 species.
Mesospio Gravier 1911a, M. moorei Gravier 1911a; Prostomium anteriorly blunt; peristomium without
only species. lateral wings. Four pairs of branchiae, either cirriform
Prostomium with frontal horns. Branchiae present or pinnate or both, first present from setiger 2. Branchial
from the second setiger, completely free of the post- parapodia with large postsetal lobes. All anterior setae
setal lobes. Notosetae all capillaries; neuropodia with capillaries; in median and posterior parapodia uncini
uncini present from setiger 15. All anal cirri similar, also present.
short and tapering.
Pseudomalacoceros Czerniavsky 18816, Nerinides can-
Microspio Mesnil 1896, Spio mecznikowianus Claparede tabra Rioja 1918; 11 species.
1870a; 10 species. Prostomium anteriorly blunt or pointed; occipital
Prostomium anteriorly rounded. Branchiae present antenna present. Branchiae present from setiger 2; at
from setiger 2; partially fused to the notopodial postsetal least partially fused to the notopodial postsetal lobes.
lobes. Notosetae all capillaries; neuropodia posteriorly Notosetae all capillaries; neurosetae include capillaries
also with uncini. and uncini.

Pseudopolydora Czemiavsky 1881 b, Polydora antennata Tripolydora Woodwick 1964, T. spinosa Woodwick
Claparede 1870a; I I species. 1964, only species.
Prostomium anteriorly blunt or bifid. Branchiae first Prostomium anteriorly rounded. Branchiae present
present from setiger 7. Fifth setiger modified with large, from setiger 2. Fifth setiger modified, with a few mod-
thick setae arranged in a horseshoe-shaped pattern. ified setae only. Notosetae all capillaries; neurosetae
Notosetae all capillaries; neurosetae include capillaries include capillaries and trifid uncini first present from
and posterior uncini. setiger 9.
Pygospio Claparede 1863, P. elegans Claperede 1863;

Taxonomic Note
3 species.
Prostomium anteriorly rounded. Branchiae concen- The above survey of the genera follows in the main
trated in a short posterior region, except in the males Foster's (1971) survey of the fauna from the Gulf of
where a single pair is present on the second setiger; all Mexico, except that I have preferred to list all the taxa
are fused to the notopodial postsetal lobes. Notosetae all at the generic level, rather than use some of them as
capillaries; neurosetae include capillaries and uncini. subgenera. From the number of species listed for each
genus, it will be noted that I have been rather mom
Rhynchospio Hartman 1936b, R. arenincola Hartman conservative than Foster in retaining species. The family
1936b; 4 species. is one of the better studied, and best understood families.
Prostomium with frontal horns. Branchiae present In addition to Foster's extensive review cited above,
from setiger 2, partially fused to the notopodial post- Hannerz (1956) and Soderstrom (1920) reviewed the
setal lobes. All notosetae capillaries; neurosetae include major taxa. The biology of the more common forms
capillaries and uncini. also has been rather well studied, especially the repro-
Scolecolepides Ehlers 1907, S. benhami Ehlers 1907; ductive biology of members of Polydora and Boccardia
3 species. has been the subject of several studies by Blake (e.g.
Blake 1971).
Prostomium with frontal horns. Branchiae present
from setiger 1, limited to the anterior part of the body.
Anterior setae all capillaries; posterior setae include Incertae Sedis
capillaries and uncini in both rami. Aberranm Hartman 1965, A. enigmatica Hartman 1965;
only species.
Scolelepis Blainville 1828, Lumbricus squamatus O.F.
Paired spioniform palps present; prostomium otherwise
Milller 1806; 20 species.
without appendages, eyes absent. Pharynx muscularized.
Prostomium usually pointed, rarely blunt. Branchiae
One apodous segment present; all others with biramous
present from setiger 2, partially fused to the notopodial
parapodia. All setae simple capillaries. Dorsal and ventral
postsetal lobes. Neuropodial postsetal lobes notched at
cirri and small branchiae present.
least in posterior setigers. Anterior setae all capillaries;
According to Hartman (1965) this differs from the true
posterior parapodia with uncini and capillaries in both
spionids in that it has a muscular, rather than saclike
rami.
pharynx; it further differs from the spionids in that it
Spio Fabricius 1785, Nereis filicornis O. F. Muller 1776; completely lacks hooks of any kind. This is not a larval
15 species. form, in that one of the specimens described was
Prostomium anteriorly rounded. Branchiae present ovigerous.
from setiger 1, at least anteriorly fused with the noto- The structure of the proboscis may not be a very im-
podial postsetal lobes. Notosetae all capillaries; neuro- portant feature in this group as noted by Orrhage (1966).
setae include capillaries and uncini. The total absence of uncini may not be particularly sur-
prising, considering how many spionid genera may lack
Spiophanes Grube 1860, S. kroeyeri Grube 1860, 16 such hooks in either of the two rami, or over long
species. stretches of the body. However, it is considered best
Prostomium blunt or with frontal horns; occipital to leave the genus as a free-standing genus, until it can
antenna present. Branchiae absent. First neuropodia be better investigated.
with stout hooks; all other anterior setae capillaries;
posterior neuropodia with uncini. Invalid Genera
Streblospio Webster 1879a, S. benedicti Webster 1879a, Aonis Audouin and Milne Edwards 1833c, see Scolelepis
2 species. Aonopsis Wagner 1885, indeterminable
Prostomium conical. A single pair of branchiae on Aricideopsis Johnson 1901, see Laonice
setiger 1. Setiger 2 with a dorsal collar between the Bilobaria Sveshnikov 1959, larval forms
notopodia. Notosetae all capillaries; neurosetae anteriorly Carazzia Mesnil 1896, see Pseudopolydora
all capillaries, posteriorly also uncini. Chaetosphaera Haecker 1896, larval forms

Cheironotus Costa 1861b, indeterminable (?Polydora)

Colobranchus Schmarda 1861, see Scolelepis
Ctenospio Sars 1867, see Prionospio
Diplotis Montagu 1815, see Polydora
Dipolydora Verrill 1881, see Polydora
Euspio McIntosh 1915, see Spio
Hekaterobranchus Buchanan 1890, see Streblospio
Heterospio Czemiavsky 1881 b, see Spio
Kinbergella McIntosh 1909, see Prionospio
Leipoceras Mobius 1874, see Polydora
Leucodora Johnston 1838, see Polydora
Mandane Kinberg 1866b, indeterminable
Neopygospio Berkeley and Berkeley 1954, see Pseudo-
polydora
Nerine Johnston 1838, see Scolelepis
Nerinides Mesnil 1896, see Pseudomalacoceros
Nerinopsis Elders 1912, larval form
Paranerine Czerniavsky 1881, we Aonides
Perialla Kinberg 1866, questionably Boccardia
Prospio Mesnil 1896, hypothetical
Protopolydora Czerniavsky 1881 b, see Polydora
Pseudoleucodore Czerniavsky 188lb, see Polydora
Pseudonerine Czemiavsky 1881b, indeterminable
Pseudonerine Augener 1926, see Scolelepis
Pteriptyches Grube 1872a, see Prionospio
Pygophyllum Schmarda 1861, indeterminable
Scolecolepis Malmgren 1867, mis-spelling of Scole-
lepis
Spione Orsted 1844, in Quatrefages 1865, indeterminable
Spionereis Sars 1853, NOMEN NUDUM
Spionides Webster and Benedict 1887, see Laonice
Uncinia Quatrefages 1865, see Scolelepis
FIGURE 6. (A), Family MAGELGNIDAE, Magelona sp., off

Santa Barbara, California, diagram of dorsal view, 25x; (B),
FAMILY MAGELONIDAE
ventral view of the above, 25x.
CUNNINGHAM AND RAMAGE 1888
Spioniforms with long, slender bodies, separated into

two regions. Prostomium - flattened and anteriorly ovate
or truncate, without appendages. Palps at the junction Invalid Genera
of the pro- and peristomium on the ventral side. Setae
Maea Johnston 1865, see Magelona
include capillaries and hooded bi- or multidentate hooks.
Papillaria Sveshnikov 1959, larval form
Magelonids currently are assigned to a single genus,
Rhynophylla Carrington 1865, see Magelona
Magelona Muller 1858, with type species M. papil-
licornis Muller 1858 and a total of about 35 described
FAMILY TROCHOCHAETIDAE PETTIBONE 1963
species. Dr. Meredith L. Jones of the Smithsonian
Institution is revising the family. It is anticipated that Body flattened cylindrical. Prostomium a flattened
more taxa may be recognized when this revision is ridge with palps present at the postectal comers. First
completed. parapodia directed forwards along the sides of the pro-
Magelonids are common in sandy bottoms; they are stomium. Setae capillaries, acicular falcate spines,
rather characteristic with the flattened, shovel-shaped fringed setae and straight spines.
prostomium often much wider than the rest of the animal, The family is recognized for a single genus, Tro-
which tends to be rather threadlike with very long seg- chochaeta Levinsen 1884 with type species Disoma
ments. Magelonids build only very flimsy tube-structures multisetosum Orsted 1844 and a total of nine recognized
and tend to move through the sediment. species. Pettibone (1963) correctly pointed out that the

name Disoma, originally applied to these animals by rami. Setae include capillaries, pectinate, plumose and
Orsted (1844), was used ten years earlier by Ehrenberg acicular setae. Dorsal and ventral cirri spindle or bottle-
for a protozoan. shaped.
Trochochaetids are long, slender non-tubicolous
spionifotms mainly reported from shallow water in soft
substrates. They appear to be nowhere truly numerous,
but frequently are reported from areas where quantita-
tive benthic investigations have been undertaken.
Invalid Genera
Cherusca Muller 1858, indeterminable
Disoma Orsted 1844, see Trochochaeta
Disomides Chamberlin 1919c, see Trochochaeta
Nevaya McIntosh 1911, see Trochochaeta
Pilearia Sveshnikov 1963, larval forms
Thaumastoma Webster and Benedict 1884, see Tro-
chochaeta
The family Kalaminochaetidae Nolte, 1941 was de-
scribed for two genera, Kalaminochaeta and Kalum-
maria both based on pelagic larval forms, possibly of
a Trochochaeta, but not identifiable. The family and
both genera are here considered invalid.
FAMILY POECILOCHAETIDAE HANNERZ 1956

Spioniforms with long, slender bodies. Prostomium
small with either a frontal or a median antenna. Palps
present at the postectal comers. First segment with or
FIGuRE 7. Family POECILOCHAETIDAE, Poecilochaetus
without setae, with one or two pairs of tentacular cirri. johnsoni, off Loon Point, Santa Barbara, California, 35 m,
Parapodia biramous, with lateral sense-organs between 25X.
Key to Genera
1. Antenna frontal, first segment with series of long setae Poecilochaetus
2. Antenna median, first segment asetigerous Elicodasia
Generic Definitions Taxonomic Notes

Elicodasia Laubier and Ramos 1973, E. mirabilis Setae are present from the second segment in Elico-
Laubier and Ramos 1973; only species. dasia and from the first in Poecilochaerus.
Poecilochaetids with a median antenna, without
nuchal organ. First segment is asetigerous and first FAMILY HETEROSPIONIDAE HARTMAN 1963a
setiger lacks dorsal and ventral cirri. Neuropodial
Body with short thoracic and greatly prolonged
acicular spines are present in most setigers from seg-
abdominal setigers. Prostomium blunt without append-
ment 4.
ages; palps attached at the postectal corners. Long,
Poecilochaetus Claparede 1875, P. fulgoris Claparede filiform branchiae on the thoracic setigers. Setae in
in Ehlers 1875; 12 species. nearly complete cinctures around the abdomen; most
Poecilochaetids with a frontal antenna, nuchal organs simple capillaries, but some neurosetae are thickened
present. First segment with long setae forming a cephalic and gently falcate.
cage and second setiger similar to all following seg- The family is known for a single genus, Heterospio
ments. Neuropodial acicular spines present in a few Ehlers 1874 with type species H. longissima Ehlers
anterior segments only. 1874 and for a total of four species.

Heterospionids have been reported from widely

scattered areas, mainly in deep shelf and slope sedi-
ments; the family is poorly known and the several
species may be considerably more common than their
scattered records indicate. They are rather fragile and
non-descript and thus easily overlooked if quantitative
studies are not undertaken.
Invalid Genus
Longosoma Hartman 1944c, see Heterospio
Suborder Chaetopteriformia
Short palps on prostomium; pharynx a simple, non-

eversible tube. Uncini present.
FAMILY CHAETOPTERIDAE MALMGREN 1867
Body with two or three distinct body-regions. Peri-

stomium may have one or two pairs of tentacular cirri;
palps of varying lengths always present. Anterior region
with uniramous parapodia, median and posterior re-
FIGURE 8. (A), Family CHAETOPTERIDAE, Chaetopterus
gions with biramous parapodia. Setae include capillary, variopedatus, modified from various sources, natural size; (B),
li mbate setae and modified spines in setiger 4. Pec- Family CIRRATULIDAE, Tharyx moniloceras, after Hartman,
tiniform uncini present in posterior setigers. 1969, 80x.
Key to Genera
Ia. Median notopodia bilobed 2

lb. Median notopodia never bilobed, may be fused 3
2a. (I a) A pair of small tentacular cirri present at the base of the large palps Phyllochaetopterus
2b. (I a) Tentacular cirri absent, palps large Spiochaetopterus
3a. (lb) Some median notopodia fused to form dorsal fans, palps very short Chaetopterus
3b. (Ib) Notopodia never fused; palps large Mesochaetopterus
Generic Definitions with uncini; posterior region with pointed notopodia

with internal acicula; neuropodia with uncini.
Chaetopterus Cuvier 1827, Tricoelia variopedatus
Renier 1804; only species. Phyllochaetopterus Grube 1863, P. gracilis Grube
Body divided in three distinct regions; anterior re- 1863; 17 species.
gion with uniramous parapodia with lancet-shaped Body with three distinct regions; antennae present.
setae, except setiger 4, which has stout spines. Median Median notopodia foliaceous, bilobed and asetigerous,
region with biramous parapodia; some posterior noto- neuropodia with pectinate uncini; posterior region with
podia fused to form dorsal fans, all notopodia of median pointed notopodia supported by internal acicula; neuro-
region asetigerous; neuropodia with pectinate uncini; podia with pectinate uncini.
posterior region with long, pointed notopodia with a
Spiochaetopterus Sars 1853, S. typicus Sars 1856; 12
few contained acicula, but no setae; neuropodia with
species.
pectiniform uncini.
Body with three distinct body-regions; antennae
Mesochaetopterus Potts 1914, M. taylori Potts 1914; absent. Median notopodia bilobed and foliaceous;
11 species. neuropodia with pectinate uncini; posterior region with
Body with three regions; antennae absent. Median pointed notopodia, supported by internal acicula, neuro-
notopodia all simple, asetigerous; median neuropodia podia with pectinate uncini.

Invalid Genera
Leptochaetopterus Berkeley 1927, see Spiochaetop-

terus
Mesotrocha Leuckart and Pagenstecher 1855, see
Phyllochaetopterus
Ranzania Claparede 1870b, see Mesochaetopterus
Ranzanides Chamberlin 1919c, see Mesochaetopterus
Telepsavus Costa 1861a, see Spiochaetopterus
Tricoelia Renier 1804, see Chaetopterus
Suborder Cirratuliformia
One pair of grooved palps, or a group of grooved

palps on one or several post-peristomial segments.
Prostomium without appendages, eversible pharynx
a thick ventral pad.
FAMILY CIRRATULIDAE CARUS 1863
Body cylindrical. Prostomium conical or blunt;

peristomium fused with at least two segments. Para- FIGURE 9. (A), Family HETEROSPIONIDAE, Heterospio
podia reduced; slender filiform orr clavate branchiae catalinensis, modified from Hartman, 1944c, 12x; (B), Family
ACROCIRRIDAE, Acrocirris fronrifilis, after Banse, 1969,
present on at least some setigers. All setae simple in- about 5x; (c), Acrocirris crassilis, lateral view, after Banse,
cluding capillaries and curved or excavate hooks. 1969, about 12x.
Key to Genera
Ia. A pair of long grooved palps attached on the anterior dorsum 2
l b. Two groups of grooved tentacular cirri present 5
2a (Ia). All setae slender, distally pointed Tharyx
2b (la). At least some setae either curved hooks or spines 3
3a (2b). Acicular setae distally excavate; body usually dark green or brown Dodecaceria
3b (2b). Acicular setae not excavate, body usually light colored 4
4a (3b). Acicular spines in posterior segments distally entire Chaetozone
4b (3b). Acicular spines in posterior segments distally bi- or multifid Caulleriella
5a (lb). All setae acicular, falcate spines Pseudocirratulus
5b (lb). At least some capillary setae present 6
6a (5b). Anterior region without long tentacular structures Raricirrus
6b (5b). One or more anterior segments with groups of long tentacular cirri or branchiae 7
7a (6b). Dorsal tentacular cirri first present posterior to the anteriormost branchiae Cirriformia
7b (6b). Dorsal tentacular cirri first present from the same segment as the anteriormost branchiae 8
8a (7b). Tentacular cirri on one segment only Cirratulus
8b (7b). Tentacular cirri on two or more segments Timarete
Generic Definitions Paired palps inserted dorsally anterior to, or at the

fast setiger. Setae include capillaries and distally entire,
Caulleriella Chamberlin 1919c, Cirratulus bioculatus curved spines.
Keferstein 1862; 16 species.
Cirratulus Lamarck 1801, Lumbricus cirratus O.F.
Paired palps inserted dorsally anterior to, or at the
Muller 1776, 25 species.
first setiger. Setae include capillaries and distally bi-
dentate or multidentate curved spines. Cirratulids without palps; tentacular cirri present on
one segment only; anteriormost branchiae present from
Chaetozone Malmgren 1867, C. setosa Malmgren 1867; the same segment. Setae include capillaries and acicular
1 9 species. spines that are usually entire distally.

Cirriformia Hartman 1936a, Terebella tentaculata Taxonomic Notes

Montagu 1808; 26 species. appears likely that the generic division of the
ItCirauldswhop;tenaculris
palpate cirratulids will change with future revisions.
on one segment only; anteriormost branchiae present
The Tharyx-complex is presently rather confused, and
anterior to the tentacular cirri. Setae include capillaries
additional characters will have to be included to clarify
and curved, distally entire spines.
the relations between the several species described in
Dodecaceria Orsted 1843b, D. concharum Orsted this genus.
1843b; 16 species.
Palps present; branchiae present on a few segments;
filiform or clavate. Anterior setigers with capillary Invalid Genera
setae; median and posterior ones also with excavate
Ambo Chamberlin 1918, see Timarete
stout spines. Posterior part of body often flattened
Archidice Kinberg 1866b, see Cirratulus
and expanded.
Audouinia Quatrefages 1865, see Cirriformia
Pseudocirratulus Augener 1922, P. kingstonensis Cirratulispio McIntosh 1911, questionably Cirratulus
Augener 1922; only species. or Chaetozone
Palps absent, cirri and branchiae present on most Cirrhineris Blainville 1818, indeterminable
segments; two achaetous anterior segments present. Heterocirrus Grube 1855, see Dodecaceria
Setae all gently curved, entire spines or hooks. Heterocirrus Saint-Joseph 1894, see Caulleriella and
in part Tharyx
Raricirrus Hartman 1961, R. maculatus Hartman 1961;
Labranda Kinberg 1866b, see Cirriformia
only species.
Mesocirrineris Czerniavsky 18816, see Caulleriella
Body anteriorly narrow, then abruptly broad at
Monticellina Laubier 1961a, see Tharyx
setiger 9 and tapering from there to the posterior end.
Naraganseta Leidy 1855, see Dodecaceria
Branchiae present on some anterior segments; setiger 13
Pentacirrus Wesenberg-Lund 1958, see Timarete
with a pair of papillae. Setae include capillaries in the
Promenia Kinberg 18666, see Cirratulus
notopodia and neuropodia and curved hooks with
dentate cutting edges.
Tharyx Webster and Benedict 1887, T. acutus Webster FAMILY ACROCIRRIDAE BANSE 1969
and Benedict 1887; 22 species.
Palps Body either slender and elongate, or short and
anterior to or at the first b
maggot-shaped. Prostomium blunt, paired palps usually
setae slender, capillary and smooth, bututst some- All
present. Several segments usually crowded near the
ti mes with ithserrated
serrated
ated cutting edges.
anterior end; these segments usually asetigerous. Para-
Timarete Kinberg 1866b, Cirratulus anchylochaetus podia biramous with small parapodia; notosetae seg-
Schmarda 1861; 8 species. mented and spinose; neurosetae compound hooded
Palps absent; tentacular cirri present on at least two falcigers.
anterior setigers; branchiae present from the same seg- The acrocinids recently were recognized as a separate
ment as the first tentacular cirri. Setae include noto- family, and even more recently, the genus Flabelligella
podial capillaries and anteriorly neuropodial capillaries; known from several deep-water locations was trans-
posterior neuroopodia with gently curved spines. ferred to this family (Orensanz 1974b).
Key to Genera
Ia. Branchiae absent Flabelligella
lb. Branchiae present 2
2a (lb). Palpal bases well separated; epithelium densely papillated Macrochaeta
2b (I b). Palpal bases nearly abutting, epithelium nearly smooth Acrocirrus
Generic Definitions the gill. Thorax of 12 segments; a row of papillae

ventral to the neurosetae in all setigers.
Acrocirrus Grube 1872b, Heterocirrus frontifilis Grube
1 860; 8 species. Flabelligella Hartman 1965, F. papillata Hartman
Acrocirrids with palps originating close together. 1965; 5 species.
Four pairs of gills; a papilla on third segment below Acrocirrids without branchiae and palps; separation

between thorax and abdomen indistinct. Body-surface difficult. However, genera and species cannot be iden-
densely papillated. Anterior segments uniramous in tified without this information. I have found it useful
some species. to rotate the specimen while counting the segments,
Macrochaeta Grube 1850, Nais clavicornis Sars 1835; trying to observe the setae as the light catches them.
6 species. The rostrate uncini look truncate in this sort of obser-
Acrocirrids with bases of palps separated by at least vation, and the capillaries will retain their tapered
the width of the palps. Usually four pairs of branchiae. appearance.
Number of thoracal segments variable; epidermis usu-
ally densely papillated. All parapodia biramous.
Invalid Genus
Ledon Webster and Benedict 1887, see Macrochaeta
ORDER CAPITELLIDA
Prostomium without appendages; palps absent. One
or two anterior asetigerous segments. Parapodia bi-
ramous; neuropodia long transverse welts in at least
some setigers. All setae simple, including capillaries
and rostrate uncini.
FAMILY CAPITELLIDAE GRUBE 1862
Body cylindrical, usually slender. Prostomium short

and truncate, without appendages. Thorax and abdomen
defined on internal structures and parapodial develop-
ment; generally, thorax with capillary setae and ab-
domen with rostrate hooded hooks.
Capitellids are among the most commonly encoun-
tered polychaetes; some forms, such as Capitella
capitata are considered pollution indicators, in that
they are capable of invading areas where disastrous
defamation of natural or man-made causes has taken
place (Grassle and Grassle 1974).
The anterior thorax of most capitellids is usually
strongly areolated and they are usually reddish pink B
or brown. In general appearance, they are perhaps the C
most earthwormlike of the common marine polychaetes. FIGURE 10. (A), Family CAPITELLIDAE, Notomactus tenuis,
after Hartman, 1947, 7x; (b), hooded hook of the above, 850x;
The characteristic rostrate uncini are small, making (C), Family ARENICOLIDAE, Arenicola, near marina. modi-
the determination of the first occurrence of these setae fied from several sources, about natural size.
Key to Genera
Ia. Three first setigers with capillary setae only 2

l b. At least four setigers with capillary setae only 4
2a (Ia). Some thoracic segments with flanged hooks; twelve thoracic segments present Peresiella
2b (Ia). Flanged hooks absent; either ten or thirteen thoracic segments present 3
3a (2b). Setigers 7-8 with genital spines; ten thoracic segments Capitomastus 9
3b (2b). Genital spines absent; 13 thoracic segments Heteromastides
4a (lb). Four first setigers with capillary setae only 5
4b (Ib). At least five anterior setigers with capillary setae 8
5a (4a). Genital spines present 6
5b (4a). Genital spines absent 7

6a (5a). Genital spines in setigers 7-8; ten thoracic segments present Capitomastus d
6b (5a). Genital spines in setigers 8-9; nine thoracic segments present Capitella (part)
7a (5b). Thorax with I I segments Mediomastus
7b (5b). Thorax with 12 segments Parheteromastus
8a (4b). Five setigers with capillary setae 9
8b (4b). At least six anterior setigers with capillary setae only 10
9a (8a). First setiger with both noto- and neuropodia; branchiae present Heteromastus
9b (8a). First setiger with notopodia only; branchiae absent Parheteromastides
l 0a (8b). Six anterior setigers with capillary setae only 11
l 0b (8b). At least seven anterior setigers with capillary setae 13
I la (l0a). Genital spines in setigers 8-9 Capitellides
1lb (l0a). Genital spines absent 12
I 2a (I1 b). Thorax with 11 segments Neomediomastus
12b (IIb). Thorax with 12 segments Barantolla
l3a (l0b). Seven anterior setigers with capillary setae only 14
13b (l0b). At least nine anterior setigers with capillary setae only 17
1 4a (13a). Genital spines present 15
14b (13a). Genital spines absent 16
15a (14a). Notopodial cirriform branchiae on some abdominal segments Branchiocapitella
15b (l4a). Branchiae absent Capitella
l6a (14b). Thorax with eight segments; all thoracic setigers with capillary setae only Leiocapitellides
l6b (14b). Thorax with 12 segments; posterior thoracic setigers with hooks Neoheteromastus
17a (13b). Nine anterior setigers with capillary setae only 18
17b (13b). Ten or more anterior setigers with capillary setae 19
18a (I 7a). One anterior asetigerous segment present; notopodial acicula absent; thorax with ten segments ...
Pseudoleiocapitella
I 8b (17a). Anterior asetigerous segment absent; notopodia with acicula; thorax with nine segments ... Pulliella
19a (l7b). Ten first setigers with capillary setae only 20
l 9b (17b). At least 11 anterior setigers with capillary setae 23
20a (19a). First setiger with both noto- and neuropodia Decamastus
20b (19a). First setiger with notopodium only 21
21a (20b). All thoracic setigers with capillary setae; all abdominal setigers with hooks only; mixed segments
absent Capitellethus
21b (20b). At least one segment with mixed hooks and capillary setae 22
22a (21b). Two first abdominal segments with mixed hooks and capillary setae; thorax with eleven seg-
ments Neonotomastus
22b (21b). Last thoracic segment with both hooks and capillary setae; twelve thoracic segments present
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Paraleiocapitella
23a (19b). Eleven anterior setigers with capillary setae only 24
23b (19b). Twelve or more anterior setigers with capillary setae only 27
24a (23a). At least some capillary setae in the first two abdominal setigers 25
24b (23a). All abdominal segments with hooks only 26
25a (24a). First setiger with neuropodia only; first two abdominal setigers with capillary setae in both
rami Notodasus
25b (24a). First setiger with both noto- and neuropodia; first two abdominal setigers with hooks in the neuro-
podia and mixed hooks and capillary setae in the notopodia Mastobranchus
26a (24b). Abdominal notopodial tori nearly coalesce; notopodial hooks absent Rashgua
26b (24b). Abdominal notopodial tori well separated; notopodial hooks present Notomastus
27a (23b). Twelve setigers with capillary setae only 28
27b (23b). Thirteen or more setigers with capillary setae only 30
28a (27a). Anal plaque with imbedded spines present Scyphoproctus
28b (27a). Anal end rounded or with anal cirri 29
29a (28b). Thirteenth setiger with both hooks and capillary setae Leiochrus
29b (28b). No setiger with both hooks and capillary setae present Leiochrides
30a (27b). Thirteen anterior setigers with capillary setae only 31
30b (27b). At least 15 anterior setigers with capillary setae 35

31a (30a). First setiger with notopodia only 32

31b (30a). First setiger with both noto- and neuropodia 33
32a (31a). Thorax with 14 or 15 segments; one segment with both capillary setae and hooks present
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Leiocapitella
32b (31 a). Thorax with 15 to 18 segments; three or four segments with both capillary setae and hooks pres-
ent Pseudocapitella
33a (31b). Thorax with 19 segments; branchiae neuropodial Capitobranchus
33b (31b). Thorax with maximally 15 segments; branchiae, if present, notopodial 34
34a (33b). All notopodial setae capillary Protomastobranchus
34b (33b). Abdominal notopodial setae hooks Dasybranchus
35a (30b). At least 16 setigers with capillary setae only; additional four segments have both setae and hooks in
the notopodia Eunotomastus
35b (30b). Thoracic count otherwise; maximally two transitional segments present 36
36a (35b). First setiger with both noto- and neuropodia Dasybranchetus
36b (35b). First setiger with notopodia only 37
37a (36b). Twenty setigers with capillary setae Lumbricomastus
37b (37b). Seventeen or 18 setigers with capillary setae only Anotomastus
Generic Definitions Thorax with 11 segments; asetigerous segment ab-

sent; first setiger with notopodia only. All thoracic
Anotomastus Hartman 1947, Eunotomastus gordioides
setigers with capillary setae only; branchiae absent.
Moore 1909; only species.
Thorax with 18 or 19 segments; one anterior asetiger- Capitellides Mesnil 1897b, C. giardi Mesnil 1897b;
ous segment present; first setiger with notopodia only. 3 species.
Up to segment 17 or 18 capillary setae only; segments Thorax with nine segments; asetigerous segment
18 and 19 transitional with notopodial setae and neuro- absent; first setiger complete. First six setigers with
podia with mixed hooks and setae. Palmately branched capillary setae only; next three with hooks, except
branchiae in posterior segments. notopodia 8-9, which has genital spines. Branchiae
absent.
Barantolla Southern 1921, B. sculpta Southern 1921;
2 species. Capitobranchus Day 1962, C. macgregori Day 1962;
Thorax with 12 segments; one anterior asetigerous only species.
segment present; first setiger complete; up to segment 7 Thorax with 19 segments; one asetigerous segment
with capillary setae only; segment 8 with mixed setae present; first setiger complete. Up to segment 15 with
and hooks in notopodia and hooks in neuropodia. capillary setae only; next four segments mixed with cap-
Branchiae may be present in posterior segments. illary setae in the notopodia and hooks in the neuropodia.
Neuropodial branchiae present in posterior setigers.
Branchiocapitella Fauvel 1932, B. singularis Fauvel
1 932; 2 species. Capitomastus Eisig 1887, Capitella minima Langerhans
Thorax with nine segments; asetigerous segment 1 881; 3 species.
absent; first setiger complete. Up to segment 7, capil- Thorax with ten segments; one asetigerous segment
lary setae only; segments 8-9 with genital spines. present; first setiger complete. Males: up to segment 5
Notopodial cirriform branchiae present. with capillary setae only, the next two segments with
hooks, segments 8-9 with genital spines. Females:
Capitella Blainville 1828, Lumbricus capitatus Fabricius
up to segment 4 with capillary setae; the next three
1780; 15 species.
segments with hooks and segments 8-9 with genital
Thorax with nine segments; asetigerous segment
spines.
absent; first setiger complete. Either four first segments
with capillary setae only; then three segments with Dasybranchetus Monro 1931, D. fauveli Monro 1931;
mixed hooks and setae in both rami, then genital spines only species.
in segments 8-9, or first seven segments with capillary Thorax with 16 segments; one asetigerous segment
setae only; then segments 8-9 with genital spines. present; first setiger complete; all thoracic setigers with
Branchiae absent. capillary setae. Branchiae not seen.
Capitellethus Chamberlin 1919c, Capitellides dispar Dasybranchus Grube 1850, Dasymallus caducus Grube
Ehlers 1907; 3 species. 1846; 10 species.

Thorax with 14 segments; one asetigerous segment Lumbricomastus Thomassin 1970, L. tulearensis Thom-
present; first setiger complete; all thoracic setigers assin 1970; only species.
with capillary setae only. Simple or composite noto- Thorax with 21 segments; one asetigerous segment
podial branchiae present. present; first setiger with notopodia only. All thoracic
setigers with capillary setae only. Branchial processes
Decamastus Hartman 1963a, D. gracilis Hartman on posterior notopodia.
1963a; 2 species.
Thorax with I I segments; one asetigerous segment Mastobranchus Eisig 1887, M. trinchesii Eisig 1887;
present; first setiger complete; all thoracic setigers with 4 species.
capillary setae only; branchiae absent. Thorax with 12 segments; one asetigerous segment
present; first setiger complete. All thoracic setigers
Eunotomastus McIntosh 1885, E. grubei McIntosh with capillary setae only; two first abdominal segments
1885; only species.
with mixed capillary setae and hooks in the notopodia
Genus poorly defined: approximately 20 thoracic
and hooks in the neuropodia. Notopodial branchiae
segments present; of which 16 have capillary setae
present.
only and the next four have mixed setae and hooks
in the notopodial fascicles. Mediomastus Hartman 1944b, M. californiensis Hart-
man 1944b, 7 species.
Heteromastides Augener 1914, H. bifidus Augener Thorax with I I segments; one asetigerous segment
1914; 2 species. present; first setiger complete. Up to segment 5 with
Thorax with 13 segments; one asetigerous segment capillary setae only; thereafter, all hooks. Branchiae
present; first setiger complete. Up to segment 4, cap-
absent.
illary setae only; segment 5 with mixed capillary setae
and hooks in both rami; from segment 6, hooks only. Neoheteromastus Hartman 1960, N. linens Hartman
Branchiae absent. 1960; only species.
Thorax with 12 segments; one asetigerous segment
Heteromastus Eisig 1887, Capitella fliformis Claparede
present; first setiger with notopodia only. Up to seg-
1 864; 7 species.
ment 8 with capillary setae only; segment 9 with capil-
lary setae in the notopodia and hooks in the neuropodia.
Branchiae not seen.
illary setae only; from segment 7, all setae hooks.
Notopodial branchiae present. Neomediomastus Hartman 1969, Mediomastus glabrus
Leiocapitella Hartman 1947, L. glabra Hartman 1947; Hartman 1960; only species.
2 species. Thorax with 11 segments; one asetigerous segment
Thorax with 14 or 15 segments; one asetigerous present; first setiger complete. Up to segment 7 with
segment present; first setiger with notopodia only. capillary setae only, then all setigers with hooks.
Up to segment 14 with capillary setae only; segment Small notopodial branchial processes present in far
15 with capillary setae in the notopodia and hooks in posterior setigers.
the neuropodia. Branchiae absent. Neonotomastus Fauchald 1972, N. glabrus Fauchald
Leiocapitellides Hartmann-Schroder 1960a, L. analis 1 972; only species.
Hartmann-Schroder 1960a; only species. Thorax with 11 segments; one asetigerous segment
Thorax with eight segments; one asetigerous segment present; first setiger with notopodia only. All thoracic
present; all thoracic segments with capillary setae setigers with capillary setae only; first abdominal noto-
only; first abdominal with notopodial capillary setae podia with capillary setae, second with mixed hooks
and neuropodial hooks. Branchiae absent. and capillary setae. First abdominal neuropodia with
mixed hooks and capillary setae; second with hooks
Leiochrides Augener 1914, L. australis Augener 1914; only. Branchiae not seen.
7 species.
Thorax with 13 segments; one asetigerous segment Notodasus Fauchald 1972, N. magnus Fauchald 1972;
present; first setiger complete. All thoracic setigers 2 species.
with capillary setae only. Branchiae not seen. Thorax with 12 segments; one asetigerous segment
present; first setiger with neuropodia only. All thoracic
Leiochrus Ehlers 1908, L. alutaceus Ehlers 1908; setigers and the first two abdominal setigers with cap-
only species. illary setae only. Branchiae not seen.
Thorax with 13 or 14 segments; one asetigerous
segment present; first setiger complete. Up to segment Notomastus Sars 1850, N. latericeus Sars 1850; 34
13, capillary setae only; segment 14 with mixed setae species.
and hooks in both rami. Branchiae absent. Thorax with 12 segments; one asetigerous segment

present; first setiger complete. All thoracic setigers First setiger complete. Branchiae absent; notopodial
with capillary setae only. Branchiae may be present. acicula present.
Paraleiocapitella Thomassin 1970, P. mossambica Rashgua Wesenberg-Lund 1949, P. rubrocincta Wesen-
Thomassin 1970; only species. berg-Lund 1949; only species.
Thorax with 12 segments; one asetigerous segment Thorax with 12 segments; one asetigerous segment
present; first setiger with notopodia only. Up to seg- present; first setiger complete; all thoracic setigers
ment 11 with capillary setae only; segment 12 with with capillary setae. Dorsal abdominal tori nearly
notopodial capillary setae and neuropodial hooks. coalesce, lack notopodial hooks. Simple branchiae
Branchiae absent. present.
Parheteromastides Hartmann-Schroder 1962a, P. mul- Scyphoproctus Gravier 1904, S. djiboutensis Gravier
tioculatus Hartmann-Schroder 1962a; only species. 1904; 7 species.
Thorax with I I segments; one asetigerous segment Thorax with 13 segments; one asetigerous segment
present; first setiger with notopodia only. Up to seg- present; first setiger complete. All thoracic setigers with
ment 6, capillary setae only; segment 7 with both cap- capillary setae only. Expanded anal plaque with acicular
illary setae and hooks and neuropodia with hooks i mbedded spines present.
only. Branchiae absent.
Parheteromastus Monro 1937a, P. tenuis Monto 1937a;
Taxonomic Notes
only species.
Thorax with 12 segments; one asetigerous segment The generic sub-division of the capitellids is un-
present. Up to segment 5, capillary setae only; all satisfactory; it is based on the number of thoracic
other segments with hooks only. Branchiae absent. segments, which may be difficult to see, and on the
distribution of the capillary and hooked setae, which
Peresiella Harmelin 1968. P. clymenoides Harmelin
is easily enough seen, but poorly understood in terms
1968; 2 species.
of variability. The key to genera follows the traditional
system, but has been based on the number of segments
present; first setiger with notopodia only. Up to seg-
with capillary setae, rather than on the number of
ment 4, capillary setae only; the remainder of thorax
thoracic segments, since the former is the more easily
with modified, flanged hooks in at least some segments,
observed character. A review only, such as the current
capillary setae and normal hooks in the others.
one, cannot solve the problem of the number of valid
Protomastobranchus Gallardo 1968, P. huloti Gallardo capitellid genera, which must be based on the vari-
1968; only species. ability of all observable characters in a large amount
Thorax with 13 or 14 segments; one asetigerous of material.
segment present; first setiger complete. All notopodia The genus Capitita Hartman 1947, is considered
with limbate capillary setae only; up to segment 13 or here a synonym of Mediomastus Hartman 1944b, as
1 4, capillary setae in neuropodia also; from there on, first suggested by Hartmann-Schroder (1962a).
hooks. The genus Bucherta Rullier (1965a) is considered
Pseudocapitella Fauvel 1913, P. incerta Fauvel 1913; as the posterior end of a capitellid, probably of the
2 species. genus Dasybranchus as first suggested by Gallardo
Thorax with 15 to 18 segments; one asetigerous (1968).
segment present; first setiger with notopodia only. Up As stated above, the generic identification of capitel-
to segment 15, capillary setae only; next three with lids is at best difficult. To make a review somewhat
notopodial capillary setae and neuropodial hooks. easier, a table has been constructed to indicate in a
Branchiae absent. different manner the relationships between the genera.
It is of the utmost importance that the two concepts,
Pseudoleiocapitella Harmelin 1964, P. fauveli Har- segments and setigers, be kept apart, since both are
melin 1964; only species. used in key features of the capitellids.
Thorax with ten segments; one asetigerous segment
ill ary setae only; first two abdominal segments with Invalid Genera
notopodial capillary setae and neuropodial hooks.
Ancistria Quatrefages 1865, see Heteromastus
Branchiae absent.
Arenia Quartrefages 1865, see Notomastus
Pulliella Fauvel 1929, P. armata Fauvel 1929; 2 Areniella Verrill 1874, questionably Heteromastus
species. Branchoscolex Schmarda 1861, see Dasybranchus
Thorax with nine segments; all with capillary setae. Bucherta Rullier 1965a, see Dasybranchus

Capitellides Ehlers 1907, see Capitellethus Isomastus Gravier 191 la, see Capitella
Dasymallus Grube 1846, see Dasybranchus Lumbriconais Orsted 1842, in Grube 1850, see Capitella
Ditrocha Sveshnikov 1959, larval forms Sandanis Kinberg 1867b, see Notomastus
Eisigella Gravier 1901, see Notomastus Valla Johnston 1865, see Capitella
TABLE 2
Genera of Capitellidae. Arranged in order of increasing number of thoracic segments present.
Column 1: No. of thoracic segments Column 5: No. of setigers with capillary setae
present Column 6: No. of setigers with mixed capillaries and
Column 2: Presence or absence of an rostrate uncini
anterior asetigerous segment Column 7: First segment complete or incomplete
Column 3: No. of thoracic setigers Column 8: Presence of branchiae and special kinds of
Column 4: Presence of genital spines setae.
Genus 1 2 3 4 5 6 7 8
Leiocapitellides 8 + 7 - 7 - compl no branchiae
Pulliella 9 9 - 9 - comp) no branchiae notacicula pres

Capitella (1) 9 9 8-9 4 3 comp) no branchiae
Capitellides 9 9 8-9 6 - compt no branchiae
Capitella (2) 9 9 8-9 7 - comp) no branchiae
Branchiocapitella 9 9 8-9 7 compi branchiae
Capitomastus 4 l0 + 9 8-9 3 - compl no branchiae

Capitomastus d 10 + 9 8-9 4 - comp) no branchiae
Pseudoleiocapitela 10 + 9 9 2 compl no branchiae
Mediomastus 11 + 10 4 compI no branchiae

Parheteromastides 11 + 10 - 5 notop. no branchiae
Neomediomastus II + 10 - 6 - compl small knobs
Capitellethus 11 - 10 - 10 - notop. no branchiae
Decamastus 11 + 10 10 - comp) ?no branchiae
Neonotomastus 11 + 10 - 10 2 notop. ?no branchiae
Peresiella 12 + 11 3 var. notop. flanged setae

Parheteromastus 12 + 11 4 - comp) no branchiae
Heteromastus 12 + 11 5 - compl branchiae
Barantolla 12 + 11 6 1 compl ? branchiae
Neoheteromastus 12 + 11 7 1 notop. ?branchiae
Paraleiocapitella 12 + 11 10 notop. no branchiae
compI
Notomastus 12 + 11 ll +_ branchiae
notop.
Rashgua 12 + 11 11 - compl no notouncini branchiae
Mastobranchus 12 + 11 11 2 compl branchiae
Notodasus 12 + Il 13 - neuro ?branchiae
Heteromastides 13 + 12 3 1 compl no branchiae

Leiochrides 13 + 12 12 - comp) ?branchiae
Scyphoproctus 13 + 12 12 - compl anal plaque
Leiochrus 13-14 + 12-13 12 1 comp) no branchiae
Protomastobranchus 13-14 + 13-14 - 13-14 - compl no notouncini
Dasybranchus 14 + 13 13 - comp) branchiae
Leiocapitella 14-15 + 13 13 1 notop. no branchiae
Pseudocapitella 15-18 + 14-17 14 3 notop. no branchiae
Dasybranchems 16 + 15 15 comp! ?branchiae
Anotomartus 18-19 + 17-18 17-18 1 nomp. branchiae
Capitobranchus 19 + 18 14 4 compl neurop. branchiae
Lumbricomastus 21 + 20 20 - notop. branchiae
The genus Eunotomastus is too incompletely known to be included in the table.

FAMILY ARENICOLIDAE JOHNSTON 1 835 With the exception of Branchiomaldane, the are-
nicolids are not easily confused with any other poly-
chaetes. The very thick, strongly areolated epidermis
Body cylindrical, separated into two or three dis- in most forms, the distinct branchial region with their
tinct regions. Prostomium without appendages. Noto- strongly tufted branchiae and the habitat, makes it easy
podia bluntly truncate, neuropodia elongated tori. to recognize the "sand-worms" from all over the world.
Notosetae capillary or limbate, neurosetae rostrate The family has been the object of very intensive studies
hooks. Branchiae present on some setigers in a median by Wells and his students. Wells (1962) and other papers
or posterior region. established the major classificatory criteria.
Key to Genera
Ia. Body slender, branchiae first present from setiger 18, or later, as thick filaments arranged with max-
imally two or three in a tuft Branchiomaldane
lb. Body thick, branchiae first present from a more anterior setiger as thick tufts of very fine filaments . 2
2a (I b). Asetose caudal end absent Arenicolides
2b(Ib). Asetose caudal end present 3
3a (2b). Neuropodia of branchial segment approach midventrally; a single pair of oesophageal sacs pres-
ent Arenicola
3b (2b). Neuropodia of branchial segments well separated; more than one pair of oesophageal sacs pres-
ent Abarenicola
Generic Definitions Invalid Genera
Abarenicola Wells 1959, Arenicola claparedii Levin- Chorizobranchus Quatrefages 1865, see Arenicola
sen 1884; 16 species and subspecies. Clymenides ClaparBde 1863, see Arenicolides
Three body-regions, including a prebranchial and Protocapitella Berkeley and Berkeley 1932, see
branchial region as well as an asetose caudal end. Branchiomaldane
Branchiae from setiger 7. Neuropodia widely separated Pteroscolex Lutken 1864, see Arenicola
in the branchial region. More than one pair of oeso- Puparia Sveshnikov 1959, larval forms
phageal sacs. Telethusae Savigny 1818, see Arenicola
Arenicola Lamarck 1801, Lumbricus marinas Linnaeus

1758; 9 species and subspecies. FAMILY MALDANIDAE MALMGREN 1867
Three body-regions, including a prebranchial and
Capitelliform polychaetes with long cylindrical
branchial region as well as an asetose caudal end.
bodies, usually truncate at one or both ends; most
Branchiae from setiger 7. Neuropodia approach mid-
species with long, cylindrical segments. Prostomium
ventrally in branchial segments. One pair of oesopha-
without appendages, with a pair of nuchal slits and a
geal sacs present.
median cephalic keel. Notopodia short and rounded;
Arenicolides Mesnil 1898, Arenicola ecaudata Johnston neuropodia elongated ton. Notosetae smooth or spinose
1835; 2 species. capillaries; neurosetae rostrate hooks, anterior modified
Two body-regions, including a prebranchial and a spines present in several forms.
branchial region. Branchiae from setiger 12-17 to the The bamboo-worms are recognized easily in that
posterior end. All neuropodia approach midventrally. for once, the popular name of the members of this
family makes sense; the long segments with the para-
Branchiomaldane Langerhans 1881, B. vincenti Langer- podia at one end, give the worms the jointed appear-
hans 1881; only species. ance of slender bamboo-shoots. However, even though
Two body-regions, including a prebranchial and easily recognized to family, they are not as easily
branchial region. Branchiae first present from setiger 18 identified to genus and species. It is necessary to have
or behind; as thick, sessile filaments, usually no more complete specimens to get them even to sub-family;
than three filaments in a group. All neuropodia widely or at least, one must have both anterior and posterior
separated. ends from the same specimen to get them safely iden-

tifed. It is here strongly suggested that identification EUCLYMENINAE: Anterior and posterior ends with
of fragments not be undertaken, except of course when plaques, anus terminal.
one can be sure about the provenance of the fragments.
Maldanids are especially common in shelf sediments.
They are all tubicolous, with mud-walled tubes, and
are usually quite large animals, often up to 20 cm in
length. Most frequently the worms are darkly red or
orange in life, often with lighter glandular fields and
neuropodial tori.
The major taxonomic revision of the maldanids was
done by Arwidsson (1907). He established the sub-
families and by and large the currently accepted genera.
Another very important study, especially on members
of the subfamily Euclumeninae, is that of Verrill (1900).
The subfamilies are defined on the development of
the anterior and posterior ends. These can be plain and
rounded, or they may form flattened discs, plaques,
or funnels. The anal plaque may have series of marginal
anal cirri or be unadorned. The cephalic plaque always
has paired nuchal slits, but is otherwise unadorned,
except that the anterior point of the prostomium may
project as a short palpode.
The subfamilies may be defined as follows:
RHODININAE: Anterior and posterior ends without
plaques; posterior setigers with numerous encircling
collars; uncini in double rows.
LUMBRICLYMENINAE: Anterior and posterior
ends without plaques; posterior segments withoutt
collars; uncini in single rows.
NICOMACHINAE: Anterior end without plaque;
anal plaque present; uncini in single rows. FIGURE 11. (A), Family MALDANIDAE, Axiothella rubro-
MALDANINAE: Both anterior and posterior ends cincta, Tomales Bay, California, intertidal, 17x; (B), posterior
with plaques; anus dorsal. end of the above, 17x.
Key to Genera
Ia. Both cephalic and anal plaques absent 2

lb. At least anal plaque present 7
2a (Ia). Rostrate uncini in double rows, posterior segments with encircling collars RHODININAE
Rhodine
2b (Ia). Rostrate uncini in single rows, posterior segments not collared LUMBRICLYMENINAE
3
3a (2b). More than 20 setigers present Praxillura
3b (2b). Nineteen setigers present 4
4a (3b). Setiger 4 with a deep encircling collar Clymenopsis
4b (3b). Setiger 4 without collar 5
5a (4b). Pygidium with anus at the end of a simple, upturned, flattened cone Lumbriclymenella
5b (4b). Pygidium otherwise 6
6a (5b). Pygidium flattened with anus dorsal Notoproctus
6b (5b). Pygidium conical with anus terminal Lumbriclymene
7a (lb). Cephalic plaque absent, anal plaque present NICHOMACHINAE 8
7b (Ib). Both cephalic and anal plaques present 10
8a (7a). Rostrate uncini in all setigers Micromaldane
8b (7a). Acicular spines in first three setigers, thereafter rostrate uncini 9
9a (8b). Anal funnel symmetrically developed Nicomache

THE POLYCHAETE WORMS 39

1976
9b (8b). Anal funnel asymmetrical, with the dorsal side reduced Petaloproctus
Anus dorsal MALDANINAE 11
l0a (7b).
l0b (7b). Anus terminal EUCLYMENINAE 15
lla (l0a). Rostrate uncini in two or three series in most setigers Sonatsa
l lb (l0a). Rostrate uncini always in single series 12
12a (I lb). First three setigers with acicular neuropodial spines Clymaldane
12b (I lb). First setiger either without neurosetae or with rostrate uncini 13
13a (12b). Cephalic keel long and high, cephalic rim shallowly notched laterally Maldane
13b (12b). Cephalic keel short and low, cephalic rim deeply incised laterally 14
14a (13b). Numerous branchial filaments on median setigers Branchioasychis
146 (136). Branchial filaments absent Asychis
15a (l0b). Series of vascular caeca cover the surface of the two last setigers Johnstonia
15b (l0b) Vascular caeca absent 16
16a (15b). Anal plaque marginally smooth or gently crenulated 17
16b (15b). Anal plaque bordered by distinct anal cirri 18
17a (16a). First setiger with notosetae only; anal plaque gently crenulated Abyssoclymene
I7b (16a). First setiger with noto- and neurosetae; anal plaque smooth Microclymene
18a (16b). All anal cirri similar in length 19
18b (16b). Midventral anal cirrus (rarely two) distinctly longer than all other anal cirri 23
19a (18a). More than 30 setigers present Macroclymene
19b (18a). Nineteen or 20 setigers present 20
20a (19b). Acicular spines in the first neuropodia 21
20b (19b). First neurosetae rostrate uncini 22
21a (20a). Setiger 4 with a deep encircling collar Clymenella
21b (20a). Setiger 4 without a collar Isocirrus
22a (20b). First setiger with notosetae only; anal plaque with numerous cirri Maldanella
22b (20b). First setiger with both noto- and neurosetae; anal plaque with a few long cirri only Clymenura
23a (18b). More than 30 setigers present 24
23b (18b). Eighteen to 20 setigers present 25
24a (23a). Setiger 4 with a deep encircling collar Macroclymenella
24b (23a). Setiger 4 without a collar Gravierella
25a (23b). Anterior neuropodia with rostrate uncini Axiothella
25b (23b). Anterior neuropodia with acicular spines 26
26a (25b). Anal plaque with two large ventral and several shorter lateral and dorsal anal cirri Proclymene
26b (25b). A single large ventral cirrus and several shorter lateral and dorsal anal cirri present 27
27a (26b). Nuchal slits short, straight and diverging anteriorly Pseudoclymene
27b (26b). Nuchal slits long, straight and parallel 28
28a (27b). Anal cone projecting well beyond the rim of the anal plaque Praxillella
28b (27b). Anal cone low and not projecting beyond the rim of the anal plaque 29
29a (28b). Apart from the long ventral anal cirrus, all other anal cirri similar in length Euclymene
29b (28b). Anal cirri of varying lengths Heteroclymene
Generic Definitions Axiothella Verrill 1900, Axiothea catenata Malmgren

1865; 12 species.
Abyssoclymene Hartman 1967, A. annularis Hartman EUCLYMENINAE. Cephalic rim incised or entire;
1967; only species. 18-20 setigers, acicular spines absent. Anal plaque
EUCLYMENINAE. Nuchal slits straight. Nineteen with long midventral and shorter lateral and dorsal anal
setigers, first with notosetae only; anal plaque nearly cirri.
smooth, gently crenulated along margin.
Asychis Kinberg 1867b, A. atlanticus Kinberg 1867b; Branchioasychis Monro 1939c, B. colmani Monro
28 species. 1939c; 3 species.
MALDANINAE. Cephalic rim with deep lateral in- MALDANINAE. Cephalic rim deeply incised later-
cisions; cephalic keel short and low. Neurosetae absent ally; cephalic keel short and low. Neurosetae absent in
in first setiger. Branchiae absent. first setiger. Numerous gill filaments on median setigers.


Clymaldane Mesnil and Fauvel 1939, C. sibogae podia. Anal plaque with one long midventral and sev-
Mesnil and Fauvel 1939; 2 species. eral shorter anal cirri. Two last setigers covered with
MALDANINAE. Cephalic rim shallowly notched. series of vascular caeca.
Keel short and very low. Acicular spines present in
anterior neuropodia. Branchiae absent. Lumbriclymene Sars 1872, L. cylindricauda Sars 1872;
5 species.
Clymenella Verrill 1873b, Clymene torquatus Leidy LUMBRICLYMENINAE. Nineteen setigers present;
1855; 7 species. nuchal slits strongly curved. Rostrate uncini in single
EUCLYMENINAE. Eighteen to 20 setigers; nuchal rows; acicular spines present. Anal cone symmetrical,
slits long and straight. Setiger 4 with deep encircling with circular cross-section.
anterior collar; anterior neuropodia with acicular spines
or strongly reduced uncini. Anal plaque with all cirri Lumbriclymenella Arwidsson 1911a, L. robusta Ar-
evenly long. widsson 1911 a; only species.
LUMBRICLYMENINAE. Nineteen setigers present;
Clymenopsis Verrill 1900, Clymene cingulata Ehlers nuchal slits strongly curved. Uncini in single rows;
1887; 2 species. acicular spines present. Anal end a simple upturned,
LUMBRICLYMENINAE. Nineteen setigers; setiger flattened cone with anus distal.
4 with deep encircling collar; nuchal slits strongly
Macroclymene Verrill 1900, Clymene producta Lewis
angled. First three neuropodia with spines.
1897; 2 species.
Clymenura Verrill 1900, Clymene cirrara Ehlers 1887; EUCLYMENINAE. More than 30 setigers; nuchal
11 species. slits long and straight. Acicular spines present in an-
EUCLYMENINAE. About 20 setigers; with large terior neuropodia. Anal plaque with numerous evenly
triangular glandular field on setiger 8. Nuchal slits long cirri.
l ong and straight. Acicular spines absent. Anal plaque
a flattened disc with few long anal cirri; anal cone Macroclymenella Augener 1926, M. stewartensis Au-
strongly projecting. gener 1926; only species.
EUCLYMENINAE. More than 30 setigers; nuchal
Euclymene Verrill 1900, Clymene oerstedii Claparede slits long and straight. Acicular spines absent. Setiger 4
1863; 27 species. with deep encircling collar. Anal plaque with a long
EUCLYMENINAE. Eighteen to 20 setigers; nuchal midventral and numerous short, slender cirri; anal
slits long and straight. Acicular spines present. Anal cone low.
plaque with one long median and numerous evenly long
anal cirri; anal cone low. Maldane Grube 1860, M. glebifex Grube 1860; 18
species.
Gravierella Fauvel 1919, G. multiannulata Fauvel MALDANINAE. Nineteen setigers; cephalic keel
1919; only species. long and high; cephalic rim notched. Acicular spines
EUCLYMENINAE. Numerous setigers, far posterior absent.
ones campanulate. Nuchal slits long and straight.
Acicular spines absent. Anal plaque with the anal cirri Maldanella McIntosh 1885, M. antarctica McIntosh
increasing in length from the dorsal side towards the 1885; 8 species.
long midventral cirrus. EUCLYMENINAE. Nineteen setigers; nuchal slits
long and straight. Neurosetae absent from first setiger;
Heteroclymene Arwidsson 1907, H. robusta Arwidsson acicular spines absent. Anal cone large; anal plaque
1907; 3 species. with all cirri evenly short.
EUCLYMENINAE. Nineteen setigers; nuchal slits
long and straight. Acicular spines in first neuropodia. Microclymene Arwidsson 1907, M. acirrata Arwidsson
Anal plaque with one long midventral and numerous 1907; 3 species.
shorter anal cirri of varying lengths. Anal cone low. EUCLYMENINAE. Nineteen to 20 setigers; nuchal
slits long and straight. Acicular spines absent. Anal
lsocirrus Arwidsson 1907, Clymene planiceps Sars plaque smooth-rimmed.
1872; 5 species.
EUCLYMENINAE. About 20 setigers; nuchal slits Micromaldane Mesnil 1897a, M. ornithochaeta Mesnil
long and straight. Acicular spines in anterior neuro- 1897a; 2 species.
podia. Anal plaque with all cirri of the same length. NICOMACFIINAE. Strongly curved, avicular uncini
on all setigers; collars absent. Anal plaque with crenu-
Johnstonia Quatrefages 1850b, J. clymenoides Quatre-
lated margin.
fages 1850b; 2 species.
EUCLYMENINAE. Twenty-two setigers; nuchal Nicomache Malmgren 1865, Sabella lumbricalis Fa-
slits long and straight. Acicular spines in the first neuro- bricius 1780; 16 species.

NICOMACHINAE. Rostrate uncini in single rows; Taxonomic Notes

acicular spines in the first three neuropodia. Anal
The genus Chaponella Rullier 1972 is considered
plaque with subequal anal cirri. here in the Sabellidae; it resembles closely members
Notoproctus Arwidsson 1907, N. oculatus Arwidsson of the genus Euchone, except for its loss of tentacular
1907; 9 species. crown.
LUMBRICLYMENINAE. Nineteen setigers, pros- The genus Sonatsa was described originally in its
tomium large, hoodlike. Acicular spines in anterior own subfamily; the posterior end of the only known
species remains unknown. The anterior end closely
neuropodia. Anal plaque flat, smooth-rimmed, with
anus dorsal. resembles well-known species of Maldane and the
genus is considered here in the subfamily Maldaninae.
Petaloproctus Quatrefages 1865, P. terricolus Qua-
t refages 1865; 11 species. Invalid Genera
NICOMACHINAE. Nineteen setigers. Acicular
spines in the first neuropodia. Anal plaque asymmetri- Arwidssonia McIntosh 1914b, see Euclymene
cal with the dorsal side strongly reduced. Axiothea Malmgren 1865, see Axiothella
Caesicirrus Arwidsson 191lb, see Euclymene
Praxillella Verrill 1881, Praxilla praetermissa Malm- Chrysothemis Kinberg 1867b, see Asychis
gren 1865; 20 species. Clymene Oken 1815, indeterminable
EUCLYMENINAE. Eighteen to 19 setigers; nuchal Clymene Savigny 1818, indeterminable
slits long and straight. Acicular spines in anterior Heteromaldane Ehlers 1908, see Maldane
neuropodia. Anal plaque short, with a long midventral Iphianissa Kinberg 1867b, questionably Praxillella
and several shorter anal cirri. Anal cone very large, Leiocephalus Quatrefages 1865, see Euclymene
projecting well beyond the rim of the anal plaque. Leiochone Grube 1868b, see Maldane and Asychis
Leiochone Saint-Joseph 1894, see Clymenura
Prarillura Verrill 1880, P. ornata Verrill 1880; 6 Maldanopsis Verrill 1900, see Asychis
species. Mandrocles Kinberg 1867, indeterminable
LUMBRICLYMENINAE. Twenty to 29 or more Neco Kinberg 1867b, see Mylitta
setigers; nuchal slits strongly curved. Acicular spines Nicomachella Levinsen 1884, see Petaloproctus
in anterior and posterior neuropodia. Paraxiothea Webster 1879a, see Clymenella
Petaloclymene Augener 1918, incompletely known
Proclymene Arwidsson 1907, Clymene muelleri Sars Praxilla Malmgren 1865, see Praxillella
1856; only species. Promaldane Mesnil 1897a, hypothetical
EUCLYMENINAE. Nineteen setigers; nuchal slits Sabaco Kinberg 1867b, see Asychis
anteriorly curved. Acicular spines present in anterior
neuropodia. Anal plaque with two large midventral
and several shorter anal cirri. ORDER OPHELIIDA
Prostomium without appendages; palps absent. Max-

Pseudoclymene Arwidsson 1907, Clymene quadrilobata
i mally one anterior asetigerous segment present. Para-
Sars 1856; 2 species.
podia uniramous or (usually) biramous. All setae
EUCLYMENINAE. Eighteen to 20 setigers, nuchal
simple, including capillary setae. Rostrate hooks ab-
slits short, straight and diverging anteriorly. Acicular
spines present; anal cirri all of the same length except sent. All neuropodia short and truncate.
Composite setae have been reported from a single
the larger midventral one.
species of the family Scalibregmidae; this form may
Rhodine Malmgren 1865, R. loveni Malmgren 1865; be incorrectly assigned to family and in some respects
8 species. resembles members of the Syllidae.
RHODININAE. Numbers of setigers variable; first
setigers with notosetae only; acicular spines absent. FAMILY OPHELIIDAE MALMGREN 1867
Numerous encircling collars on posterior setigers.
Uncini in double rows. Body with a limited number of segments; often with
a deep ventral furrow. Prostomium without appendages,
Sonatsa Chamberlin 1919c, S. meridionalis Chamber- blunt or conical. Parapodia biramous or uniramous,
lin 1919c; only species. with small button-shaped parapodial lobes; all setae
MALDANINAE. Numbers of setigers unknown; capillary, either smooth or marginally dentate.
prostomium with high, long cephalic keel; cephalic Opheliids have a series of three very distinct body-
rim notched. Rostrate uncini in two or three series in shapes. Some are short, thick and grub-shaped; others
some setigers. are very slender, nearly smooth and torpedo-shaped.

Finally, some are anteriorly inflated and posteriorly in sandy and muddy bottoms and have been studied
cylindrical or narrow. Opheliids are common animals extensively by ecologists interested in sandy beaches.
Key to Genera
la. Body fusiform without ventral groove 2
lb. Body cylindrical with at least posterior ventrum deeply grooved 4
2a (I a). Branchiae absent Kesun
2b (la). Branchiae present 3
3a (2b). First setiger appear in front of the mouth; all setae smooth Travisia
3b (2b). First setiger appear behind the mouth; some setae spinose Dindymenides
4a (lb). Ventral groove present in posterior part of body only 5
4b (1b). Ventral groove present along the whole body 6
5a (4a). Three body-regions, including inflated head, inflated anterior part and narrow posterior part;
branchiae in posterior region only Euzonus
5b (4a). Body not clearly regionated, inflated anteriorly and grooved posteriorly; branchiae from setigers
8-10, if present Ophelia
6a (4b). Branchiae absent 7
6b (4b). Branchiae present 8
7a (6a). Lateral eyes absent Tachytrypane
7b (6a). Lateral eyes present Polyophthalmus
8a (6b). Lateral eyes present Armandia
8b (6b). Lateral eyes absent 9
9a (8b). Anal tube short, all anal cirri of similar length Antiobactrum
9b (8b). Anal tube long, two internally attached ventral cirri present, dorsal anal cirri short 10
l0a (9b). Branchiae, if present, along the whole body Ophelina
l0b (9b). Branchiae always present, limited to the posterior end of body only Ammotrypanella
Generic Definitions
Ammotrypanella McIntosh 1879, A. arctica McIntosh
1879; only species.
Ventral groove present along the whole body;
branchiae present and limited to the posterior end only.
Anal tube long and narrow, with two internally attached
ventral cirri. Lateral eyes absent.
Antiobactrum Chamberlin 1919c, Ophelina brasiliensis

Hansen 1882; only species.
Ventral groove present along the whole body;
branchiae present. Anal tube short and with all anal
cirri of the same length. Lateral eyes absent; expanded
palpode at the tip of the prostomium.
Armandia Filippi 1861, A. cirrhosa Filippi 1861; 18
species.
Ventral groove along the whole body; branchiae
present. Lateral eyes present; anal tube long and slen-
der, with paired long internally attached ventral cirri
and shorter dorsal cirri.
Dindymenides Chamberlin 1919c, Dindymene concinna
Kinberg 1866b; 3 species.
Body short and grublike without ventral furrow.
FIGURE 12. (A), Family OPHELIIDAE, Ophelia rathkei, modi-
fied from Hartmann-Schroder, 1971, about 5x; (B), Family
Branchiae present. First setiger appears behind the SCALIBREGMIDAE, Scalibregma injlatum, off Anacapa
mouth; some setae spinose. Island, California, 90 m, 5x.

Euzonus Grube 1866, E. arcticus Grube 1866; 10 Taxonomic Notes

species. I am presently following Hartmann-Schreder (1971)
Body with three regions; a head consisting of pros- and others in accepting Ophelina as the valid name for
tomium and one or two setigers, an anterior swollen
worms often referred to under the generic name Am-
region and a posterior region with ventral groove.
motrypane. Hartmann-Schroder (1971) also recognized
Branchiae limited to posterior region. Last setiger of
a series of subfamilies within this family. There are
anterior region with specialized notopodia.
two distinct groups of genera within the family, but
the differences between the groups and the size of the
Kesun Chamberlin 1919c, K. fusus Chamberlin 1919c;
family does not seem to warrant such treatment.
5 species.
Body short and grublike without ventral furrow. Invalid Genera
Branchiae absent. Last setigers without epipodial pads;
anal cylinder furrowed, but without papillae. Aloysina Claparede 1864, see Polyophthalmus
Ammotrypane Rathke 1843, see Ophelina
Ophelia Savigny 1818, O. bicornis Savigny 1818; 32 Cassandane Kinberg 1866b, see Ophelia
species. Dindymene Kinberg 1866b, see Dindymenides
Body fusiform with inflated anterior end and ventral Ladice Kinberg 1866b, see Ophelina
furrow posteriorly. Branchiae rarely absent, on most Nais Dujardin 1839, see Polyophthalmus
setigers from setigers 8-10. No notopodial lobes Neomeris Costa 1844, see Ophelia
modified. Nitetis Kinberg 1866b, see Ophelia
Omaria Grube 1869b, see Ophelina
Ophelina Orsted 1843b, O. acuminata Orsted 1843b; Pygophelia Sars 1869, indeterminable
44 species. Terpsichore Kinberg 1866b, see Ophelina
Body with ventral furrow along the whole length. Urosiphon Chamberlin 1919c, see Ophelina
Branchiae presently, rarely absent, on most setigers.
Anal cone long, with paired internally attached ventral FAMILY SCALIBREGMIDAE MALMGREN 1867
c irri and shorter dorsal cirri. Lateral eyes absent. Body short and stocky or long and slender, often
Polyophthalmus Quatrefages 1850a, Nais pitta Du- anteriorly inflated. Prostomium anteriorly bifid or with
T-shaped frontal horns. Parapodia biramous, with
jardin 1839; 3 species.
Body with ventral furrow along the whole length. small, triangular or button-shaped parapodia. Setae
Branchiae absent; lateral eyes present. Anal tube short include capillaries and furcate setae and in some cases
acicular spines. Composite setae reported in one species.
with small anal cirri.
The scalibregmids contained up to a t;.w years ago
Tachyrrypane McIntosh 1879, T. jeffreysii McIntosh a well-circumscribed group of forms; all charae.erized
1879; only species. by having capillary and furcate setae; usually combined
Body with ventral furrow along the whole length. with a strongly wrinkled or areolated epidermis. During
Branchiae and lateral eyes absent. Anal tube cut away last ten years several new forms have been added,
ventrally forming an open hood with marginal anal making a definition of the family more difficult. Some
cirri. of these forms (Proscalibregma and Scalispinigera)
may in fact not be as closely associated with this family
Travisia Johnston 1840, T. forbesii Johnston 1840, as previously indicated; especially the latter resembles
20 species. forms usually associated with the order Phyllodocida.
Body short and grublike without a ventral groove. No attempts have been made below to move these
Branchiae present; posterior setigers with epipodial genera to other families, but it is anticipated that the
pads. Anal cirri all short and thick. All setae smooth; content of this family may change rather drastically
first setiger appears before the mouth. in the near future.
Key to Genera
Ia. Body short, maggotlike and inflated 2
lb. Body long, sometimes anteriorly inflated, but always with a slender posterior end 4
2a (Ia). Two or three anterior asetigerous segments Neolipobranchius
2b (la). One anterior asetigerous segment 3
3a (2b). Branchiae present Polyphysia
3b (2b). Branchiae absent Lipobranchius

4a (Ib). Neurosetae composite heterogomph spinigers Scalispinigera

4b (Ib). Neurosetae capillary, furcate or acicular 5
5a (4b). Furcate setae absent Proscalibregma
5b (4b). Furcate setae present 6
6a (5b). Median parapodia with long, digitate postsetal lobes Scalibregmides
6b (5b). Postsetal lobes short and triangular or absent 7
7a (6b). Prostomium with long, cirriform frontal antenna Scalibregmella
7b (6b). Prostomium T-shaped, without appendages 8
8a (7b). Neither dorsal nor ventral cirri present 9
8b (7b). At least ventral cirri present I1
9a (8a). Acicular setae present Asclerocheilus
9b (8a). Acicular setae absent 10
10a (9b). Eyes present Hyboscolex
l0b (9b). Eyes absent Kebuita
I I a (8b). Branchiae present 12
11b (8b). Branchiae absent 14
12a(1Ia). Acicular spines absent Scalibregma
12b (I la). Acicular spines present 13
13a (12b). Eyes present Parasclerocheilus
13b (12b). Eyes absent Sclerobregma
14a (IIb). Dorsal cirri absent Sclerocheilus
I 4b (11 b). Dorsal cirri present Pseudoscalibregma
Generic Definitions Parasclerocheilus Fauvel 1928b, P. branchiatus Fauvel

Asclerocheilus Ashworth 1901, Lipobranchius inter- 1928b; 2 species.
medius Saint-Joseph 1894; 4 species. Body elongated; prostomium T-shaped; eyes present.
Body elongated, prostomium T-shaped, eyes absent. One asetigerous segment present. Branchiae present.
One asetigerous segment. Branchiae, dorsal and ventral Dorsal cirri absent, ventral cirri present in posterior
cirri absent. Acicular spines in up to three anterior setigers. Acicular spines present in up to four anterior
setigers; otherwise with furcate and capillary setae. setigers; other setae include capillaries and furcate
setae.
Hyboscolex Schmarda 1861, H. longiseta Schmarda
1861; 3 species. Polyphysia Quatrefages 1865, Eumenia crassa Orsted
Body elongated, prostomium T-shaped. Branchiae, 1943b; 2 species.
dorsal and ventral cirri absent. Setae include capillaries Body short and maggotlike; prostomium bilobed;
and furcate setae. Eyes present. one asetigerous segment present. Branchiae present.
Acicular spines absent; setae include furcate and capil-
Kebuita Chamberlin 1919c, Eumenia glabra Ehlers lary setae.
1887; 2 species.
Body elongated, prostomium T-shaped. Branchiae, Proscalibregma Hartman 1967, P. linea Hartman 1967;
dorsal and ventral cirri absent. Setae include capillaries only species.
and furcate setae. Eyes absent. Body elongated; prostomium trapezoidal; eyes ab-
sent. No asetigerous segment present; all setae capil-
Lipobranchius Cunningham and Ramage 1888, Eu-
lary; anus within a collar-shaped pygidium.
menia jeffreysii McIntosh 1869; only species.
Body short and maggotlike, prostomium bilobed; one Pseudoscalibregma Ashworth 1901, Scalibregma parva
asetigerous segment present. Branchiae absent. Aci- Hansen 1878; 3 species.
cular setae absent; furcate and capillary setae present. Body elongated; prostomium T-shaped. One asetiger-
ous segment present. Dorsal and ventral cirri present
Neolipobranchius Hartman and Fauchald 1971, N.
in posterior setigers. Branchiae absent. Furcate and
glabrus Hartman and Fauchald 1971; only species.
capillary setae present; acicular spines present in some
Body short and maggotlike; prostomium a conical
forms.
lobe. Two or three asetigerous anterior segments pres-
ent. Acicular spines, furcate setae, branchiae and dorsal Scalibregma Rathke 1843, S. inflatum Rathke 1843,
and ventral cirri absent. Eyes absent. 4 species.

Body elongated; prostomium T-shaped. Eyes absent. ORDER PHYLLODOCIDA

One asetigerous segment present. Branchiae present.
Prostomium usually with at least one pair of anten-
Capillary and furcate setae present; acicular spines
nae; palps, if present, frontal or frontolateral. Max-
absent.
imally two pairs of jaws present, but jaws often absent.
Scalibregmella Hartman and Fauchald 1971, S. an- Eversible pharynx always muscular and cylindrical.
tennata Hartman and Fauchald 1971, only species. Parapodia distinct in most forms, supported by acicula
Body elongated; prostomium quadrangular with cir- at least in one ramus.
riform long antennae and nuchal organs. Dorsal and Apart from the amphinomids and eunicids and as-
ventral cirri absent; smooth capillary setae in both sociated families, this order contains most members
rami, furcate setae in notopodia only. A muscular of the old order ERRANTIA. It is by far the most
eversible pharynx present. speciose of the polychaete orders. For that reason, it
was considered appropriate to erect suborders and
Scalibregmides Hartmann-Schroder 1965, S. chilensis
superfamilies within the order, even if some families
Hartmann-Schroder 1965; only species.
are left without specific intermediate designation.
Body elongated, prostomium T-shaped. One asetiger-
Most members of this order are modified moderately
ous segment present. Branchiae absent. Parapodia with
from the assumed ancestral condition (Fauchald 1974a)
long, slender postsetal lobes in median and posterior
in that they have biramous parapodia and only one, or
setigers. Acicular setae absent, furcate and capillary
perhaps a few segments involved in the anterior tagma.
setae present.
Several families show varying levels of cephalization,
Scalispinigera Hartman 1967, S. oculata Hartman indicating that this process is going on independently
1967, 2 species. in these taxa. Similarly, a change in function of the
Body elongated; prostomium rectangular; eyes pres- notopodia from locomotory to defensive appears to
ent. One asetigerous segment present. Notosetae all have taken place independently in several of the groups.
capillary; neurosetae all composite heterogomph spin- It is possible to divide this order into several orders
igers. Branchiae, furcate setae, dorsal and ventral recognizing as such, taxa characterized as suborders
cirri absent. here. However, the Phyllodocida is characterized by
the same characters as all the other orders of polychaetes
Sclerobregma Hartman 1965, S. branchiata Hartman
whereas the suborders of Phyllodocida are characterized
1965; only species.
by a slightly different set of characters, making such a
Body elongated; prostomium T-shaped, one asetiger-
change undesirable.
ous segment present. Eyes present. Branchiae present.
Acicular spines in first setiger; capillary and furcate
setae present. Suborder Phyllodociformia
Sclerocheilus Grube 1863, S. minutus Grube 1863; Phyllodocida with at least two pairs of prostomial
5 species. appendages and at least two pairs of tentacular cirri.
Body elongated, prostomium T-shaped; one asetiger- Palps absent; eversible pharynx unarmed; first para-
ous segment present. Branchiae absent; ventral cirri podia lateral.
present on posterior setigers; dorsal cirri absent. Acic-
ular spines, furcate and capillary setae present.
FAMILY PHYLLODOCIDAE WILLIAMS 1851
Taxonomic Note Phyllodociformia with long and slender bodies;

prostomium with four or five antennae, eyes, when
The genus Oncoscolex Schmarda 1861, is considered present, small. Two to four pairs of tentacular cirri
here a synonym of Hyboscolex Schmarda as suggested present. Parapodia uniramous (usually) or biramous
by Day (1967). with the notopodia represented by a short stalk and
l arge, foliose dorsal cirri held erect over the dorsum,
rarely with acicula or setae. All neurosetae composite;
Invalid Genera
notosetae, when present, simple.
Eumenia Orsted 1843b, see Polyphysia The phyllodocids are common shallow-water poly-
Eumeniopsis Bidenkap 1895, see Polyphysia chaetes, more commonly associated with hard sub-
Eusclerocheilus Hartman 1967, see Pseudoscalibregma strates than with sands and muds. They are frequently
Gwasitoa Chamberlin 1919c, see Kebuita highly colored in life, and these colors are diagnostic,
Oligobranchus Sars 1846, see Scalibregma but fade very rapidly upon preservation. The phyl-
Oncoscolex Schmarda 1861, see Hyboscolex lodocida produce copious amounts of mucus when
FtcuaE 13. (A), Family ALCIOPIDAE, Vanadis formosa, off central California, pelagic, lOx; (B), seta of the above, 112x;
(C), Family PHYLLODOCIDAE, Anaitides groenlandica, Cance Bay, Alaska, 43-70 m, 12x; (D), seta of the above, 100x;
(E), median parapodium of the above, 19x; (F), Family LOPADORHYNCHIDAE, Lopadorhynchas errans, central Pacific Ocean,
deep pelagic, seta from median setiger, 125x; (G), anterior end of the above, lOx; (H), Family PONTODORIDAE, Pontodora
pelagica, modified after Uschakov, 1972, about 50x.

disturbed, and must be relaxed carefully before preser- a stereo microscope, and I have found it helpful to use
vation. If possible, each specimen should be preserved a low light-angle, so that strong shadows fall across
separately. Important recent revisions include Hartmann- the specimen.
Schroder (1971); Uschakov (1972) and Banse (1973). In certain groups of species, the ventral tentacular
cirri (on the second segment) may be foliose, often
Important identificatory characters and methods of assymmetrical and flattened. Distortions in fixation
observations: of ordinary tapering ventral tentacular cirri may be
i
The most important characters in the Phyllodocidae misleading. The flattening of the ventral c rri is very
includes the number and arrangement of antennae on striking and once seen, this character is never again
the prostomium and the numbers of tentacular cirri confused with the distortions one can find in preserved
and their arrangement on the first few segments (Berg- material.
strom 1914). This latter character may be somewhat The papillation of the eversible pharynx is another
troublesome, in that varying fusions obscure the prim- important character and if the pharynx is not everted,
itive arrangement. In principle, the first segment has a dissection may be necessary. This is easily done, either
single, dorsal pair of tentacular cirri; the next segment ventrally or dorsally on most specimens, but may be
usually both a dorsal and a ventral pair, and the third difficult to do on smaller animals. However, in most
segment has dorsally a tentacular citrus and ventrally cases it cannot be avoided.
a normal parapodium. As stated above, discrepancies Biramous and uniramous parapodia in the phyllo-
from this pattern are common, and are of great im- docida are exceedingly similar, and the difference
portance. essentially is defined by convention: the parapodia
The best way to observe these structures is to hold are considered biramous if the notopodia, i.e. the stems
the specimen with a pair of forceps so that one can of the dorsal cirri, contain internal acicula, or if setae
look in at the animal in three-quarter dorsal view, and are present. It may be necessary to make parapodial
then rotate the animal slowly from dorsal to lateral preparations and observe these under the compound
positions. In this manner it is possible to follow the microsocpe to ascertain the presence or absence of the
segmental furrows, most easily visible laterally, over internal acicula. The blood-vessels in the notopodial
towards the dorsal side, and reductions and fusions rudiments may resemble the internal acicula, so the
can be assessed. This whole process is best done under observations have to be rather careful.
Key to Genera
la, Two antennae present Cirrodoce

lb. At least four antennae present 2
2a (Ib). Two pairs of tentacular cirri on one segment 3
2b (1b). At least three pairs of tentacular cirri on two or three segments 4
3a (2a). Eversible pharynx diffusely papillated or smooth Eteone
3b (2a). Eversible pharynx with large lateral papillae in rows Mysta
4a (2b). Three pairs of tentacular cirri on two or three segments 5
4b (2b). Four pairs of tentacular cirri on three segments 7
5a (4a). Third segment with dorsal cirri Lugia
5b (4a). Third segment without dorsal cirri 6
6a (5b). Second segment with setae; dorsal and ventral cirri broadly ovate Mystides
6b (5b). Second segment without setae; dorsal and ventral cirri lanceolate Hesionura
7a (4b). Parapodia biramous (with notacicula and sometimes with notosetae) 8
7b (4b). Parapodia uniramous (without notacicula or setae) 11
8a (7a). Ventral tentacular cirrus (on segment 2) foliaceous Hesperophyllum
8b (7a). Ventral tentacular cirri digitate or subulate 9
9a (8b). First tentacular segment a complete ring Austrophyllum
9b (8b). First tentacular segment dorsally reduced 10
l0a (9b). First segment with two pairs of cirri and setae Nipponophyllum
IOb (9b). First segment with a single pair of cirri; setae first present from the second segment .. Notophyllum
Ila (7b). Five antenna present 12
11b (7b). Four antennae present 22
12a (I la). Ventral tentacular cirri (on segment 2) foliose 13

12b (I Ia). Ventral tentacular cirri digitate or subulate 16

13a (12a). First segment dorsally reduced 14
13b (12a). All tentacular segments complete rings 15
14a (13a). Second tentacular segment with setae Sige
I4b (13a). Second tentacular segment without setae Pterocirrus
15a (13b). Second tentacular segment with setae; proboscis papillated Steggoa
15b (13b). Second tentacular segment without setae; proboscis smooth Notalia
16a (12b). Nuchal epaulettes present 17
16b (12b). Nuchal epaulettes absent 18
17a (16a). First and second tentacular segment free from each other and complete rings Vitiazia
17b (16a). First and second tentacular segment fused; first segment dorsally reduced Vitiaziphyllum
18a (16b). First and second tentacular segment fused; pharynx smooth Eumida
I8b (16b). All three tentacular segments free from one another; pharynx papillated 19
19a (18b). Reniform ventral cirri broadly attached transverse to the long axis of the parapodia 20
19b (18b). Ventral cirri narrowly attached; usually narrow and elongated 21
20a (19a). Pharyngeal papillae distally in six rows Bergstroemia
20b (19a). Pharyngeal papillae dispersed Clavadoce
21a (196). Pharynx with few papillae; second segment with setae Pirakia
21b (19b). Pharynx with numerous papillae; second segment without setae Eulalia
22a (I Ib). All three tentacular segments form complete rings
. . ........ Protomystides and Pseudeulalia (these two cannot be separated on current information).
22b (l lb). First two segments fused dorsally 23
23a (22b). Nuchal papillae absent 24
23b (22b). Nuchal papillae present 25
24a (23a). Tentacular cirri flattened Nereiphylla
24b (23a). Tentacular cirri digitate or subulate Generyllis
25a (23b). First and second segment fused to each other and to the prostomium 26
256 (23b). First and second segment fused to each other at least dorsally, but free from the prostomium .... 27
26a (25a). Simple enlarged setae in segments 2-4 Chaetoparia
26b (25a). Segments 2-4 with composite spinigers Prochaetoparia
27a (25b). First segment with a pair of small dorsolateral papillae in addition to the tentacular cirri
. . . . . . . ................................................................ Prophyllodoce
27b (25b). First segment with tentacular cirri only 28
28a (27b). First to third pair of tentacular cirri and the antenna globose, four pairs of cirri digitate
. . . .....................................................................Sphaerodoce
28b (27b). All pairs of tentacular cirri and the antennae digitate or subulate 29
29a (28b). Nuchal papilla on a posterior prolongation from the prostomium Paranoids
29b (28b). Prostomium heart-shaped with the nuchal papilla in a posterior incision 30
30a (29b). Pharyngeal papillae basally in rows Anaitides
30b (29b). Pharyngeal papillae basally dispersed Phyllodoce
Taxonomic Notes Generic Definitions

The system indicated above, follows in the main the Anaitides Czerniavsky 1882, Phyllodoce groenlandica
principles first promulgated by Bergstrom (1914) and Orsted 1843a; 28 species.
followed by most later authors. It incorporates the Phyllodocids with four antennae; heart-shaped pros-
main features of the systems organized by Uschakov tomium with nuchal papilla in the crevice between the
(1972) and Banse (1973). Certain taxa here considered lobes. First segment dorsally reduced and partially
genera often are considered as subgenera. This creates fused to the second; all tentacular cirri cylindrical.
a problem of setting the relative taxonomic value of Papillae in rows on the base of the eversible pharynx.
the same character in different parts of the family, to Parapodia uniramous.
the subgeneric level in some cases and to the generic
level in others (cf. Banse 1973). For the sake of con- Austrophyllum Bergstrom 1914, Eulalia charcoti Grav-
sistency, these characters are here all considered to ier 1911b; 5 species.
be of generic value. Phyllodocids with five antennae; pentagonal or

/976 THE POLYCHAETE WORMS 49
rounded prostomium and without a nuchal papilla. All Phyllodocids with five antennae; pentagonal pros-
anterior segments complete and free from one another; tomium and indistinct nuchal papilla. First tentacular
all tentacular cirri cylindrical. Parapodia biramous. segment dorsally reduced and fused to the second; all
tentacular cirri cylindrical. Eversible pharynx smooth
Bergstroemia Banse 1973, Eulalia nigrimaculata
or with very small papillae. Parapodia uniramous.
Moore 1909; only species.
Phyllodocids with five antennae, pentagonal pros- Genetyllis Malmgren 1865, G. lutea Malmgren 1865;
tomium and without a nuchal papilla. All anterior 11 species.
segments free from one another, but the first dorsally Phyllodocids with four antennae; short wide pros-
reduced; all tentacular cirri cylindrical. Eversible tomium and no nuchal papilla. First and second ten-
pharynx with papillae in six rows distally. Parapodia tacular segments are fused and reduced dorsally; all
uniramous, ventral cirri large and reniform and attached tentacular cirri cylindrical. Eversible pharynx diffusely
transverse to the long axis of the parapodium. papillated. Parapodia uniramous.
Chaetoparia Malmgren 1867, C. nilssoni Malmgren Hesionura Hartmann-Schroder 1858, H. fragilis
1867; only species. Hartmann-Schroder 1958; 9 species.
Phyllodocids with four antennae and a nuchal papilla; Phyllodocids with four antennae, prolonged pros-
prostomium fused to the first segment, which, in turn, tomium and no nuchal papilla. Two tentacular seg-
is fused to the second. All tentacular cirri cylindrical. ments, free from each other and from the prostomium;
Specialized, enlarged simple setae in segments 2-4. all three pairs of tentacular cirri cylindrical. Third
segment without dorsal cirrus.
Cirrodoce Hartman and Fauchald 1971, C. cristata
Hartman and Fauchald 1971; only species. Hesperophyllum Chamberlin 1919a, H. tectum Cham-
Phyllodocids with two antennae, quadrangular pros- berlin 1919a; only species.
tomium and without nuchal organs. Three pairs of Phyllodocids with five antennae, short and wide
tentacular cirri on two segments; first segment with prostomium and no nuchal papilla. All tentacular
normal setae. A pair of auricular lobes attached be- segments complete; ventral cirrus foliaceous; all others
tween prostomium and first parapodia. cylindrical. Parapodia biramous; ventral cirrus large
and transversely attached.
Clavadoce Hartman 1936c, C. splendida Hartman
1936c; 2 species. Lugia Quatrefages 1865, Eteone aurantiaca Schmarda
Phyllodocids with five antennae, quadrangular pros- 1861; 3 species.
tomium and no nuchal papilla. First segment dorsally Phyllodocids with five antennae of which the median
reduced, but free from the other tentacular segments; is very small; long prostomium and no nuchal papilla.
all four pairs of tentacular cirri club-shaped and slightly Two tentacular segments with three pairs of tentacular
flattened. Eversible pharynx with diffuse papillation. cirri; all cylindrical. Third segment with dorsal cirrus.
Parapodia uniramous, ventral cirri large and reniform Parapodia uniramous.
and attached transverse to the long axis of the para-
Mysta Malmgren 1865, M. barbata Malmgren 1865;
podium.
6 species.
Eteone Savigny 1818, Nereis flava Fabricius 1780; Phyllodocids with four antennae; pentagonal pros-
40 species. tomium and no nuchal papilla. Two pairs of tentacular
Phyllodocids with four antennae; triangular or trap- cirri on first segment; all cirri cylindrical. Eversible
ezoidal prostomium and a small nuchal papilla. All pharynx with large lateral papillae in rows.
tentacular segments complete and free from one an-
Mystides Theel 1879, M. borealis Theel 1879; 8
other; two pairs of cylindrical tentacular cirri present.
species.
Eversible pharynx diffusely papillated or smooth.
Phyllodocids with five antennae; long prostomium
Parapodia un iramous.
and no nuchal papillae. Three pairs of cylindrical
Eulalia Savigny 1818, Nereis viridis Linnaeus 1767; tentacular cirri on two segments. Third segment with-
50 species. out dorsal cirri.
Phyllodocids with five antennae; pentagonal pros-
Nereiphylla Blainville 1828, N. paretti Blainville
tomium and no distinct nuchal papilla. All three ten-
1828; 3 species.
tacular segments free from one another and from the
Phyllodocids with four antennae; short wide pros-
prostomium; all tentacular cirri cylindrical. Eversible
tomium and no nuchal papilla. First and second segment
pharynx diffusely papillated; parapodia uniramous.
fused and dorsally reduced; all tentacular cirri flattened.
Eumida Malmgren 1865, Eulalia sanguinea Orsted Eversible pharynx diffusely papillated. Parapodia
1843b; 13 species. uniramous; ventral citrus reniform.

Nipponophyllum Imajima and Hartman 1964, Noto- Phyllodocids with four antennae; heart-shaped pros-
phyllum japonicum Marenzeller 1879; 3 species. tomium with a nuchal papillae between the lobes.
Phyllodocids with five antennae; rounded pentagonal First and second segment fused; first segment dorsally
prostomium and no nuchal papilla. Four pairs of reduced; all tentacular cirri cylindrical. First segment
cylindrical tentacular cirri on two segments; first seg- with a pair of small papillae dorsal to the tentacular
ment dorsally reduced, but with setae. Eversible cirri. Prostomium with diffuse papillae. Parapodia
pharynx closely papillated. Parapodia biramous. uniramous.
Notalia Bergstrom 1914, Eulalia picta Kinberg 1866b, Protomystides Czemiavsky 1882, Mystides bidentata
only species. Langerhans 1880; only species.
Phyllodocids with five antennae; pentagonal pros- Phyllodocids with four antennae and no nuchal
tomium and no nuchal papilla. All tentacular segments papilla. All tentacular segments complete rings; three
free from one another and from the prostomium; ventral or four pairs of cylindrical tentacular cirri on three
cirri foliose; all others cylindrical. Eversible pharynx segments.
smooth; parapodia uniramous, setae present from Pseudeulalia Eliason 1962, P. exigua Eliason 1962;
second segment. only species.
Notophyllum Orsted 1843b, Phyllodoce foliosa Sars Phyllodocids with four antennae and no nuchal
1835; 6 species. papilla. All tentacular segments are separate and com-
Phyllodocids with five antennae; broadly transverse plete rings; all tentacular cirri are cylindrical. Para-
prostomium and nuchal epaulettes present. All ten- podia uniramous.
tacular segments free, but first or first and second Pterocirrus Claparede 1868, Phyllodoce (Eulalia)
dorsally reduced; all tentacular cirri cylindrical. Para- macroceros Grube 1860; 10 species.
podia biramous; no setae in first segment. Phyllodocids with five antennae; pentagonal pros-
Paranaitis Southern 1914, Anaitis wahlbergi Malm- tomium and no nuchal papilla. All tentacular segments
gren 1865; 15 species. free from one another; first segment dorsally reduced;
Phyllodocids with four antennae; rounded pros- ventral tentacular cirrus foliose, all others cylindrical.
tomium and a nuchal papilla present. First and second Parapodia uniramous; setae first present in second
segment fused and dorsally reduced; all tentacular segment.
cirri cylindrical. Eversible pharynx with rows of lateral Sige Malmgren, 1865, Sige fusigera Malmgren, 1865;
papillae. Parapodia uniramous. about five species.
Phyllodoce Savigny 1818, P. laminosa Savigny 1818; Phyllodocids with five antennae, pentagonal pros-
48 species. tomium and no nuchal papilla. All tentacular segments
Phyllodocids with four antennae; heart-shaped pros- free from one another; first segment dorsally reduced;
tomium with nuchal papilla in the crevice between the ventral tentacular cirrus foliose, all others cylindrical.
lobes. First segment dorsally reduced and fused to the Parapodia uniramous, setae first present in the third
second; all tentacular cirri cylindrical. Eversible pharynx segment.
with diffuse papillation. Parapodia uniramous. Sphaerodoce Bergstrom 1914, Phillodoce quadraticeps
Pirakia Bergstrom 1914, Phyllodoce (Eulalia) punctifera Grube 1878, only species.
Gmbe 1860; 4 species. Phyllodocids with four antennae; heart-shaped pros-
Phyllodocids with five antennae; pentagonal prostomium with a nuchal papilla between the lobes. First
tomium and no nuchal papilla. All three anterior seg- and second segments fused and dorsally reduced; first
ments free from each other; all tentacular cirri cylindrical. three pairs of tentacular cirri and the antennae globose;
Eversible pharynx with few scattered papillae. Para- last pair of tentacular cirri (on third segment) digitate.
podia uniramous; setae present from second segment. Steggoa Bergstrom 1914, Eulalia magalaensis Kinberg
Prochaetoparia Bergstrom 1914, Genetyllis brevis 18666; 13 species.
Ehlers 1901; only species. Phyllodocids with five antennae, pentagonal pros-
Phyllodocids with four antennae; broadly trans- tomium and no nuchal papilla. All three tentacular
verse prostomium; a nuchal papilla present. First and segments complete rings, free from one another. Ven-
second segment fused to each other and to the pros- tral tentacular cirrus foliose; all others cylindrical.
tomium; all tentacular cirri cylindrical. Parapodia uni- Eversible pharynx diffusely papillated. Parapodia
ramous, with normal composite setae in all setigers. uniramous.
Prophyllodoce Hartman 1966b, P. hawaila Hartman Vitiazia Uschakov 1953, V. dogieli Uschakov 1953;
1966b; only species. only species.

Phyllodocids with four large and one small median Hypoeulalia Bergstrom 1914, see Eulalia
antenna; broadly truncate prostomium and paired nuchal Kinbergia Quatrefages 1865, indeterminable
epaulettes. All tentacular segments fully developed Lepadorhynchus Schmarda 1861, see Anaitides
and separate from one another; tentacular cirri cylin- Macrophyllum Schmarda 1861, see Notophyllum
drical. Parapodia uniramous. Mesoeulalia Czemiavsky 1882, indeterminable
Mesomystides Czemiavsky 1882, see Mystides
Vitiaziphyllum Uschakov 1972, V. nuchalum Uschakov
Myriana Savigny 1820, indeterminable
1972; only species.
Myriacyclum Gmbe 1880, see Eulalia
Phyllodocids with five antennae, broadly transverse
Nothis Pruvot 1885, hypothetical
prostomium and paired nuchal epaulettes. First and
Paraeulalia Czemiavsky 1882, seeNotophyllum
second segment fused; first segment reduced dorsally;
Phyllodoce Ranzani 1817, see Polyodontes (family
all tentacular cirri cylindrical. Parapodia uniramous.
Polyodontidae)
Porroa Quatrefages 1865, see Eulalia
Protocarobia Czemiavsky 1882, see Phyllodoce
Invalid Genera Pseudonotophyllum Czemiavsky 1882, see Notophyllum
Trachelophyllum Levinsen 1883, see Notophyllum
Carobia Quatrefages 1865, see Nereiphylla
Carobia (Paracarobia) Czemiavsky 1882, see Anaitides
Carobia (Protocarobia) Czemiavsky 1882, see Phyllo-
doce FAMILY ALCIOPIDAE EHLERS 1864
Cirraria Sveshnikov 1959, larval forms
Eracia Quatrefages 1865, see Eulalia and Sige Phyllodociforms with slender, transparent long bodies
Eteonella McIntosh 1874, see Eteone found exclusively pelagically. Prostomium with five
Eteonides Hartmann-Schroder 1960a, see Hesionura antennae and a pair of very large, spherical eyes. Three
Eulalides Czemiavsky 1882, see Eumida (?) to five pairs of tentacular cirri. Parapodia uniramous,
Eumenia Quatrefages 1865, error for Eunomia, inde- dorsal and ventral cirri foliaceous; setae simple or
terminable composite.
Eumidia Verrill 1873b, error for Eumida Alciopids are delicate, slender pelagic organisms,
Eunornia Risso 1826, indeterminable mainly known for their very l arge, complex camera-
Eunotophyllum Czemiavsky 1882, see Notophyllum type eyes (Hermans and Eakin 1974). The pelagic
Globiodoce Bergstrom 1914, lapsus calami for Sphaero- polychaetes recently were reviewed by Dales and
doce Peter (1972). The system indicated below, follows
Hypereteone Bergstrom 1914, see Eteone these two authors. The key to genera has been rewritten
Hypocirrus Giard 1913, indeterminable from Uschakov (1972).
Key to Genera
Ia. Several anterior segments with rudimentary parapodia 2
lb. Anterior segments with fully developed parapodia 4
2a (Ia). All setae simple capillaries Naiades
2b (la). Setae composite spinigers 3
3a (2b). Parapodia distally with a cirriform appendage Vanadis
3b (2b). Parapodia distally without a cirriform appendage Torrea
4a (1b). Parapodia with two distal cirriform appendages, nearly all setae composite spinigers Alciopa
4b (Ib). Parapodia without cirriform appendages, or with a single such appendage; simple or acicular setae
present 5
5a (4b). All setae simple 6
5b (4b). At least some composite setae present 7
6a (5a). Parapodia with a cirriform appendage Krohnia
6b (5a). Parapodia without ciriiform appendages Alciopina
7a (5b). Parapodia with citriform appendages 8
7b (5b). Parapodia without cirriform appendages Plotoheimis
8a (7b). Acicula barely extending beyond the tip of the parapodia Rhynchonerella
8b (7b). Aciculum prolonged, extending well beyond the tip of the parapodia Watelio

Generic Definitions Alciopids with rudimentary parapodia on the two

first segments. Three pairs of tentacular cirri. Para-
Alciopa Audouin and Milne Edwards 1833b, A. podia without digitiform distal cirri. All setae com-
reynaudi Audouin and Milne Edwards 1833b; one, posite spinigers with long narrow appendages.
possibly three species.
Alciopids with parapodia fully developed on the Vanadis Claparede 1870b, V. formosa Claparede
anterior segments. Each parapodium with two digiti- 1870b; 13 species.
form cirri distally. Three or four pairs of tentacular Alciopids with several segments with rudimentary
cirri. Setae include numerous composite spinigers, parapodia, maximally ten segments. Three or four pairs
but a few slender capillaries may be present. of tentacular cirri. Each parapodium with a digitiform
distal cirrus. All setae composite spinigers with long,
Alciopina Claparede and Panceri 1867, A. parasitica slender appendages.
Claparede and Panceri 1867; 2 species.
Alciopids with parapodia fully developed on anterior Watelio Stop-Bowitz 1948, Callizona gravieri Benham
segments. Parapodia without digitiform cirri. Five 1 929; only species.
pairs of tentacular cirri. All setae simple, including Alciopids with parapodia fully developed on all
numerous slender capillaries and several shorter acicular segments. Four pairs of tentacular cirri present. Each
setae. parapodium with a digitiform distal cirrus; acicula
extending well beyond the tip of the parapodia. Setae
Krohnia Quatrefages 1865, Alciopa lepidota Krohn include numerous composite spinigers with long,
1845; 2 species. slender appendages and a few simple capillaries.
Alciopids with parapodia fully developed on anterior
segments. Parapodia with a single digitiform cirrus
distally. Five pairs of tentacular cirri. All setae simple, Invalid Genera
including capillaries and thicker, acicular setae in
anterior segments and ventrally in other parapodia. Asterope Claparede 1870b, see Torrea
Callizona Greeff 1876, see Rhynchonerella
Naiades delle Chiaje 1830, N. cantrainii delle Chiaje Callizonella Apstein 1891, see Krohnia
1830; only species. Cleta Claparede 1870b, lapsus calami for Vanadis
Alciopids with rudimentary parapodia on first three Corynocephalus Levinsen 1885, see Alciopina
body segments. Three pairs of tentacular cirri. Para- Greela McIntosh 1885, see Alciopa
podia without digitiform cirrus. All setae simple Halodora Greeff 1876, see Alciopa
capillaries. Liocapa Costa 1862, see Torrea
Mauita Chamberlin 1919c, see Vanadis
Plotohelmis Chamberlin 1919c, Rhynchonerella cap-
Nauphanta Greeff 1876, see Alciopa
itata Greeff 1876; 3 species.
Alciopids with parapodia fully developed on anterior
segments. Four or five pairs of tentacular cirri present. FAMILY LOPADORHYNCHIDAE CLAPAREDE 1 868
Parapodia without digitiform distal cirrus. Setae of two
kinds including simple acicular setae and composite Exclusively pelagic, short-bodied phyllodociform
setae with slender pointed appendages. polychaetes. Prostomium with four antennae and a pair
of small eyes. Two or three pairs of tentacular cirri
Rhynchonerella Costa 1862, R. gracilis Costa 1862; present. Parapodia Imitations, dorsal and ventral cirri
5 species. cirriform; all setae composite except for modified
Alciopids with parapodia fully developed on anterior anterior setae in some forms.
segments. Five pairs of tentacular cirri present. Para- Members of this family often are considered as
podia with single digitiform distal cirri. Setae of several truly phyllodocids and are treated as such. It is pref-
kinds, including slender composite setae with long erable to segregate them in their own family; inclusion
appendages, and shorter acicular setae simple or com- in the phyllodocids would make that family very dif-
posite. ficult to define. Dales and Peter (1972) and Uschakov
Torrea Quatrefages 1865, Alciopa candida delle Chiaje (1972) have given recent reviews; but the latter con-
1 828; 2 species. sidered the lopadorhynchids as part of the phyllodocids.
Key to Genera
Ia. Setae present at the base of the tentacular cirri 2

lb. Setae absent at the base of the tentacular cirri 3
2a (la). Two pairs of tentacular cirri on one segment; parapodial cirri long and digitate Pelagobia

2b (I a). Three pairs of tentacular cirri on two segments; parapodial cirri short Maupasia
3a (Ib). First two or three parapodia modified and with large, simple setae Lopadorhynchus
3b (lb). All parapodia similar, all setae composite spinigers Pedinosoma
Generic Definitions Mastigethus Chamberlin 1919c, see Lopadorhynchus

Reibischia Bergstrom 1914, see Lopadorhynchus
Lopadorhynchus Grube 1855, L. brevis Grube 1855,
9 species.
Lopadorhynchids with three pairs of tentacular cirri FAMILY PONTODORIDAE BERGSTROM 1914
on two segments; setae absent from tentacular - seg-
Phyllodociform, small, slender, pelagic polychaetes.
ments. Two or sometimes three anterior setigers en-
Prostomium with four antennae, eyes small. Two pairs
l arged with markedly modified parapodia and simple
of tentacular cirri present. Slender, elongated uniramous
setae. Normal segments with conical setal lobes and
parapodia with long dorsal cirri; setae composite.
rounded presetal lobes. Normal setae composite with
The family is known for a single genus, Pontodora
broad, oar-shaped appendages with dentate cutting
Greeff 1879 and a single species P. pelagica Greeff
margin.
1879. It has so far been reported from warm waters
Maupasia Viguier 1886, M. caeca Viguier 1886, 4 in the Atlantic Ocean and from colder waters in the
species. Pacific Ocean (Uschakov 1972).
Lopadorhynchids with three pairs of tentacular cirri
on two segments; setae present on tentacular segments. Invalid Genus
All parapodia similar; each conical with short dorsal
Epitoka Treadwell 1943, see Pontodora
and ventral cirri. Setae include composite spinigers
with slender appendages and sometimes simple setae
Suborder Aphroditiformia
in the first setiger.
Pedinosoma Reibisch 1895, P. curtain Reibisch 1895, Phyllodocida with up to three antennae (may be
absent); one pair of tentacular cirri; long ventral palps.
only species.
Eversible pharynx, if armed, with two or four jaws.
Lopadorhynchids with two pairs of tentacular cirri
First parapodia directed forwards.
on the first segment; no setae on tentacular segment.
All parapodia similar, with small rounded dorsal cirri
Superfamily Aphroditacea
and long, slender ventral cirri. All setae composite
spinigers. Dorsal cirri alternating with elytrae in at least part
of the body; notosetae cylindrical in cross-section,
Pelagobia Greeff 1879, P. longicirrata Greeff 1879,
slender or thick. Prostomium not fused to the first
2 species.
setiger.
Lopadorhynchids with two pairs of tentacular cirri
on first segment; setae present on tentacular segment.
All parapodia similar with long setal lobes and very FAMILY APHRODITIDAE MALMGREN 1867
long dorsal and ventral cirri. All setae composite with
narrowly oar-shaped, marginally dentate appendage. Aphroditaceans with dorsoventrally flattened bodies
and one antenna present. Eversible pharynx with a pair
Invalid Genera of jaws or unarmed. Elytrae alternating with dorsal
cirri; each marginally smooth or irregularly lobed.
Haliplanella Treadwell 1943, see Maupasia Notosetae of several kinds, including in some genera
Haliplanes Reibisch 1895, see Maupasia finely attenuated felt setae and in others harpoon-
Halyplanes Reibisch 1893, see Maupasia shaped, erect, flattened spines or cylindrical spines.
Hydrophanes Claparede 1870b, see Lopadorhynchus All setae simple.
Key to Genera
Ia. Dorsum covered with felted notosetae 2
lb. Dorsum without felt, or with a very few felt setae 4
2a (Ia). Stout supportive notosetae absent; facial tubercle absent Heteraphrodita
2b (Ia). Stout supportive notosetae present; facial tubercle present 3
3a (2b). Neurosetae distally bifid Tricertia
3b (2b). Neurosetae distally entire, but sometimes subdistally spurred Aphrodita
FIGURE 14. Family APHRODITIDAE, Aphrodita refulgida, Halfmoon Bay, California, felt removed on right side, 2.5x.
4a (I b). Harpoon-shaped notosetae present Laetmonice

4b (l b). Harpoon-shaped notosetae absent 5
5a (4b). Notosetae sabre-shaped and smooth Aphrogenia
Sb (4b). Notosetae, if flattened, marginally serrated 6
6a (5b). Notosetae flattened and marginally serrated Pontogenia
6b (Sb). Notosetae cylindrical and smooth Hermionopsis
Generic Definitions
Aphrodita Linnaeus 1758, A. aculeata Linnaeus 1761; Facial tubercle absent; felt present. Stout supportive
26 species. notosetae absent.
Facial tubercle present; felt present. Harpoon-shaped
notosetae absent; other long supportive notosetae Laetmonice Kinberg 1855, L. filicornis Kinberg 1855;
present. 28 species.
Facial tubercle present; felt absent. Harpoon-shaped
Aphrogenia Kinberg 1855, A. alba Kinberg 1855; notosetae present.
5 species.
Facial tubercle present; felt absent. Harpoon-shaped Pontogenia Claparede 1868, Hermione chrysocoma
notosetae absent; notosetae long, sabrelike or smooth Baird 1865; only species.
and capillary. Facial tubercle present; felt poorly developed or
Hermionopsis Seidler 1923, H. levisetosa Seidler 1923; absent. Protective notosetae flattened and marginally
only species. serrated.
Facial tubercle present; felt absent. Harpoon-shaped Tricertia Haswell 1883, T. araeoceras Haswell 1883;
notosetae absent; notosetae long, straight and cylin- only species.
drical without asperities. Facial tubercle present; felt present. Long protective
Heteraphrodita Pettibone 1966a, H. altoni Pettibone notosetae present, flattened and curved. Neurosetae
1966a; only species. distally bifid.

Taxonomic Notes Halithea Savigny 1818, see Laetmonice

Halogenia Horst 1916b, see Laetmonice
The above treatment follows Pettibone (1966a) closely
Hermione Blainville 1828, see Laetmonice
in that several genera (e.g. Hermonia Hartman 1959),
Hermonia Hartman 1959, see Laetmonice
generally considered valid, have been excluded from
Laetmatonice ExAucToRE, see Laetmonice
the named genera. The validity of Tricertia and Hermi-
Letmonicella Roule 1898, see Laetmonice
onopsis is doubtful; however, both have been defined
Milnesia Quatrefages 1865, see Aphrodita
on characters usually used in the group, and therefore,
have been included.
FAMILY POLYNOIDAE MALMGREN 1867
Invalid Genera
Aphroditaceans with dorsoventrally flattened bodies;
Barbularia Sveshnikov 1959, NOMEN NUDUm one, two or, usually, three antennae present. Eversible
Cyanippa Castelnau 1842, see Aphrodita pharynx with four jaws. Marginally smooth or fringed
a
FIGURE 15. (A), Family
POLYNOIDAE, Halosydna brevisetosa, Sunset Bay, Oregon, intertidal, elytrae removed on left side,
lox; (B), Family POLYODONTIDAE, Polyodontes sp., off Santa Barbara, California, 84 m, elytrae removed on the right side,
15x.

elytrae alternate with dorsal cirri at least on the anterior below. They differ from Harmothoe SENSU Srstcru,
end. All setae simple, notosetae variously developed, in exactly the characters that usually are considered
but never as felt- or harpoon-setae, and usually dis- valid generic characters elsewhere in the family and
tinctly lateral in position. cannot be related to Harmothoe by overlap.
The polynoids are the most commonly occurring If possible, when preserving scale-worms, especially
scale-worms. They are usually medium-sized, rarely members of HARMOTHOINAE, care must be taken
large worms. Forms with smooth elytrae are often that scales are maintained with the specimens, since
commensals with other organisms; those with strongly these furnish one of the best diagnostic characters for
ornamented elytrae tend to be free-living. Nearly 100 species identifications. The best way of doing this, is
genera of scale-worms presently are recognized and either to bulk-relax the whole sample (MS 222, or
more are being described every year; most genera are MgCl,) or, if the specimens are larger, preserve each
well defined and the species are recognizable, but scale-worm separately.
within several of the larger genera (Harmothoe, There are no recent reviews of the whole family;
Lepidonotus, Halosydna) problems occur. Several recent authors interested in these worms, include
taxa that usually are considered as subgenera under Hartman and Hartmann-Schroder, and especially Marian
Harmothoe, have been considered as separate genera H. Pettibone of the Smithsonian Institution.
Key to Genera
la. Lateral antennae inserted posteriorly on the prostomium Cervilia
lb. Lateral antennae inserted anteriorly 2
2a (Ib). At least two antennae well developed 3
2b (lb). Median antenna well developed; lateral antennae reduced or absent MACELLICEPHALINAE
5
3a (2a). Median antenna present 4
3b (2a). Median antenna absent IPHIONINAE 7
4a (3a). Lateral antennae attached sub-distally or ventrally on the prostomium (ceratophores below the anterior
points of the prostomium which maybe produced into cephalic peaks) HARMOTHOINAE 8
4b (3a). Lateral antennae attached distally on the prostomium (ceratophores continuations of the pros-
tomium) LEPIDONOTINAE 51
5a (2b). Fifteen pairs of elytrae Macelloides
5b (2b). Maximally 13 pairs of elytrae 6
6a (5b). Prostomium consisting of two inflated lobes, 16-17 segments present Macellicephaloides
6b (5b). Prostomium a single unified lobe, with paired rounded anterior projections, 17-19 segments
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ..................................Macellicephala
7a (3b). Notosetae coarser than neurosetae Bylgides
7b (3b). Notosetae much thinner than neurosetae Iphione
8a (4a). First setigers with simple, strongly curved hooks 9
8b (4a). Anterior hooks absent 10
9a (8a). Two segments with anterior hooks Australaugeneria
9b (8a). Three segments with anterior hooks Uncopolynoe
l0a (8b). Fourteen or fewer pairs of elytrae 11
lob (8b). Fifteen or more pairs of elytrae 15
Ila(l0a). Notosetae absent Polynoella
llb (l0a). Notosetae present 12
12a (I Ib). Eight or nine pairs of elytrae present, parapodia prolonged Herdmanella
12b (I Ib). Twelve to 14 pairs of elytrae present, parapodia short 13
13a (12b). Some notocirri with spherical subdistal inflations Andresia
13b (12b). Notocirri maximally gently inflated 14
1 4a (13b). Eyes very large and confluent Robertianella
l4b (13b). Eyes small Antinoana
15a (lob). Fifteen pairs of elytrae 16
15b (l0b). Sixteen or more pairs of elytrae 42
16a (15a). More than 50 segments present, only anterior part of body covered by elytrae 17
16b (15a). Less than 45 segments present; elytrae cover the whole body, or at most, the last 10-15 segments
uncovered 23

17a (16a). Neurosetae with semilunar pockets Adyte

17b (16a). Neurosetae spinose or smooth, without pockets 18
18a (17b). Notosetae absent Heteropolynoe
18b (17b). Notosetae present 19
19a (18b). All notosetae thicker than or at least as thick as the neurosetae 20
l 9b (18b). At least some capillary notosetae present 21
20a (19a). Notosetae held erect over body Hermadion
lob (19a). Notosetae short and lateral in position Hemilepidia
21a (19b). Eyes reduced or absent, prostomium quadrangular Nemidia
21b (19b). Eyes distinct, prostomium rounded 22
22a (21b). Notosetae mainly capillary, neurosetae unidentate Enipo
22b (21b). Notosetae mainly thick and blunt-tipped, neurosetae bidentate, except in far posterior setigers
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Polynoe
23a (16b). Notosetae distinctly slenderer than the neurosetae 24
23b (16b). At least some notosetae as thick as or thicker than the neurosetae 27
24a (23a). Ventral surface with paired foliose appendages Phyllohartmania
24b (23a). Ventral surface smooth 25
25a (24b). Most neurosetae bidentate Arcteobia
25b (24b). All neurosetae unidentate 26
26a (25b). Neurosetae slender, straight spines Hartmania
26b (25b). Neurosetae distally falcate Gattyana
27a (23b). Ventral cirri foliose, ventrum papillose Phyllosheila
27b (23b). Ventral cirri digitate, ventrum usually smooth 28
28a (27b). Neurosetae in part distally trifid Gorekia
28b (27b). Neurosetae distally uni- or bidentate 29
29a (28b). Neurosetae with semilunar pockets Paradyte
29b (28b). Neurosetae spinose or nearly smooth 30
30a (29b). Notosetae with distal tufts of long hairs Barrukia
30b (29b). Notosetae without distal tufts of hairs 31
31a (30b). Neurosetae distally with capillary tips 32
31b (30b). Neurosetae distally falcate, uni- or bi-dentate 33
32a (31 a). Some neurosetae with fine, filamentous tips Antinoella
32b (31 a). All neurosetae with slender tips, but without filaments Antinoe
33a (3lb). All neurosetae unidentate 34
33b (31b). At least some neurosetae bidentate 37
34a (33a). Dorsal cirri alternating between short and long; short dorsal cirri basally inflated .... Kermadecella
34b (33a). All dorsal cirri similar in length, none basally inflated 35
35a (34b). Notosetae of two kinds, thick and slender Hesperonoe
35b (34b). All notosetae thick and similar in shape 36
36a (35b). Eyes absent Intoshella
36b (35b). Eyes present Eunoe
37a (33b). Neurosetae distally pencillate Austrolaenilla
37b (33b). Neurosetae never pencillate 38
38a (37b). Both noto- and neurosetae slender; notosetae with capillary tips Tenonia
38b (37b). Notosetae and most neurosetae coarse 39
39a (38b). Neurosetae include furcate spines with subequal tips and slender, dentate capillaries Melaenis
39b (38b). Neuropodial capillary setae absent 40
40a (39b). Some neurosetae deeply cleft with both teeth long and narrow Eucranta
4ob (39b). Neurosetae with main fangs very much larger than secondary teeth 41
41 a (4ob). Posterior eight to ten segments not covered by elytrae Lagisca
41b (40b). Whole body covered by elytrae Harmothoe
42a (I5b). Sixteen pairs of elytrae 43
42b (15b). Eighteen or more pairs of elytrae 45
43a (42a). Neurosetae with semilunar pockets Subadyte
43b (42a). Neurosetae slender and serrate, semilunar pockets absent 44
44a (43b). Notosetae distinctly serrated Leucia
44b (43b). Notosetae with a few spines Scalisetosus

45a (42b). Eighteen pairs of elytrae Acanthicolepis

45b (42b). Nineteen or more pairs of elytrae 46
46a (45b). Notosetae distinctly slenderer than neurosetae 47
46b (45b). Notosetae as thick as, or thicker than neurosetae 49
47a (46a). Neurosetae with semilunar pockets Pottsiscalisetosus
47b (46a). Semilunar pockets absent 48
48a (47b). Neuropodia with subacicular digitiform lobes Parahololepidella
48b (47b). Neuropodia with large, supra-acicular presetal lobes Grubeopolynoe
49a (46b). All neurosetae unidentate Polyeunoa
50a (49b). Neurosetae with long bare tips beyond the serrated regions Hololepidella
50b (49b). Neurosetae with very short bare tips beyond the serrated region Neohololepidella
51a (4b). Palps represented by two small tubercles Bouchiria
51b (4b). Palps well developed 52
52a (51b). Ventral cirrus heavily fimbriated, ventrum with transverse rows of three papillae on each seg-
ment Lepidofimbria
52b (51b). Ventral cirrus and ventrum smooth 53
53a (52b). Neurosetae distally bifid with several small accessory teeth in the crotch Lepidogyra
53b (52b). Neurosetae unidentate or bifid, rarely trifid; if bifid, then without accessory teeth 54
54a (53b). Prostomium anteriorly produced into two large lobes with small lateral antennae attached distally
Allmaniella
54b (53b). Prostomium anteriorly tapering into the ceratophores 55
55a (54b). Renal papillae greatly prolonged in some median setigers Bathymoorea
55b (54b). Renal papillae of similar length in all setigers 56
56a (55b). Notopodia completely reduced, neuropodium prolonged with the ventral cirrus attached near the
middle Frennia
56b (55b). Notopodia usually present; if absent, then neuropodia short with ventral cirri basal 57
57a (56b). Twelve or 13 pairs of elytrae 58
57b (56b). More than 13 pairs of elytrae 66
58a (57a). With branchial filaments on the elytrophores 59
58b (57a). Branchial filaments absent 61
59a (58a). With pseudelytrae on non-elytrophoral, alternating segments posteriorly Euphionella
59b (58a). Pseudelytrae absent 60
60a (59b). Neurosetae coarsely spinose Chaetacanthus
60b (59b). Neurosetae finely plumose Euphione
61a (58b). Notopodia absent Drieschia
61b (58b). Notopodia present 62
62a (61 b). With one very large bidentate hook in the middle of each neuropodial fascicle Sheila
62b (61b). All neurosetae similar in thickness, none bidentate hooks 63
63a (62b). Neurosetae distally trifurcate Hermenia
63b (62b). Neurosetae distally entire 64
64a (63b). Notosetae of two kinds, lancet-shaped and tapering and serrated Thormora
64b (63b). Notosetae all of one kind, usually slender and serrated 65
65a (64b). Pseudelytrae on non-elytrophoral segments Dilepidonotus
65b (64b). Pseudelytrae absent Lepidonotus
66a (57b). Fourteen pairs of elytrae 67
66b (57b). Fifteen or more pairs of elytrae 68
67a (66a). Neurosetae bidentate Lucopia
67b (66a). Neurosetae entire Podarmus
68a (66b). Only anterior half of body covered by elytrae Pseudopolynoe
68b (66b). Elytrae on most parts of the body 69
69a (68b). Fifteen pairs of elytrae 70
69b (68b). Sixteen or more pairs of elytrae 74
70a (69a). Notosetae thin Parahalosydna
70b (69a). Notosetae as thick as, or thicker than the neurosetae 71
71a (70b). All neurosetae unidentate Eulagisca

72a (7lb). Dorsal cirri of two kinds, short and expanded or long and gently inflated Malmgreniella
72b (71 b). All dorsal cirri long and slightly inflated subdistally 73
73a (72b). Notosetae nearly smooth Malmgrenia
73b (72b). Notosetae densely serrated Paralepidonotus
74a (69b). With 16 pairs of elytrae Arctonoella
74b (69b). At least 17 pairs of elytrae 75
75a (746). Seventeen to 20 pairs of elytrae 76
75b (74b). At least 21 pairs of elytrae 80
76a (75a). Notosetae absent 77
76b (75a). Notosetae present 79
77a (76a). Dorsum with a median pustule on each segment Weberia
77b (76a). Dorsal pustules absent 78
78a (77b). Neurosetae with capillary tips Drieschella
78b (77b). Neurosetae with falcate tips Alentiana
79a (76b). Notosetae numerous, lateral antennae terminal Halosydna
79b (76b). Only few notosetae present; lateral antennae subterminal Alentia
80a (75b). Notosetae present 81
80b (75b). Notosetae absent 90
81a (80a). Parapodia distally with series of long papillae Halosydnopsis
81b (80a). Parapodia distally without papillae 82
82a (81b). Neurosetae with semilunar pockets Hololepida
82b (81b). Neurosetae spinose or smooth 83
83a (82b). Notosetae thinner than most neurosetae 84
83b (82b). Notosetae as thick as, or thicker than the neurosetae 86
84a (83a). Superior neurosetae slender and spinose, inferior ones acicular Pseudohalosynda
84b (83a). All neurosetae stout 85
85a (84b). At least some neurosetae bidentate Halosydnella
85b (84b). All neurosetae unidentate Acholoe
86a (83b). All neurosetae unidentate 87
87a (86a). Ventrum with large segmentally arranged lamellae Gastrolepidia
87b (86a). Ventrum smooth Lepidastheniella
88a (86b). Lateral antennae subterminal, neurosetae few Arctonoe
88b (866). Lateral antennae terminal, neurosetae numerous 89
89a (88b). Twenty-one or 22 pairs of elytrae; neurosetae with long, curved tips Hyperhalosydna
89b (88b). Numerous pairs of elytrae; neurosetae with short, straight or curved tips Lepidasthenia
90a (8ob). Ventral cirri with two-three knobs Perolepis
90b (8ob). Ventral cirri smooth 91
9la (90b). Antennal scales present 92
91b (90b). Antennal scales absent 93
92a (91a). Setae flattened, marginally serrated with long, slender tips; up to 30 pairs of elytrae Admetella
92b (91 a). Setae flattened, finely dentate and very abruptly tapering; 23 pairs of elytrae Bathyadmetella
93a (91b). Neurosetae stout, falcate; pre- and postsetal neuropodial lobes of similar length ... Benhamipolynoe
93b (91 b). Neurosetae slender; presetal lobes distinctly longer than the postsetal ones in the neuropodia
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Telolepidasthenia
Generic Definitions LEPIDONOTINAE. Numerous segments and elytrae.

Acanthicolepis McIntosh 1900, Polynoe asperrima Cirriferous segments with T-shaped papillae. Notosetae
Sars 1861; 3 species. thinner than neurosetae, with transverse rows of spines.
HARMOTHOINAE. Eighteen pairs of elytrae, ap- Neurosetae few, unidentate with serrated subdistal
prox. 40 segments. Notosetae thicker than neurosetae, areas.
with whorls of teeth. Neurosetae bidentate with slender Admetella McIntosh 1885, A. longipedata McIntosh
secondary tooth; serrated. Dorsal cirri and antennae 1885; 2 species.
papillated. LEPIDONOTINAE. Up to 30 pairs of elytrae; 75
Acholoe Claparede 1870b, Nereis squamosa delle segments. Antennal scales at base of lateral antennae.
Chiaje 1828; only species. Notosetae absent; neurosetae long, flattened, finely

marginally serrated with smooth tips. Neuropodia setae with rows of spines. Neurosetae distally with
prolonged. slender, filamentous tips, often pilose.
Adyte Saint-Joseph 1899, Hermadion assimile McIntosh Arcteobia Annenkova 1937, Eupolynoe anticostiensis
1876; only species. McIntosh 1874; 2 species.
HARMOTHOINAE. Fifteen pairs of elytrae; long HARMOTHOINAE. Fifteen pairs of elytrae; 36 seg-
posterior region without scales. Notosetae at least as ments. Notosetae slenderer than neurosetae, with
thick as neurosetae, smooth with few spines. Neuro- blunt or capillary tips. Upper neurosetae sharp-tipped;
setae with semilunar pockets and faintly bifid tips; others bidentate.
serrations faint. Presetal neuropodial lobes longer than
Arctonoe Chamberlin 1920, Polynoe vittata Grube
postsetal ones. 1855; 4 species.
Alentia Malmgren 1865, Polynoe gelatinosa Sars LEPIDONOTINAE. Forty or more pairs of elytrae,
1835; 3 species. numerous segments. Lateral antennae subterminal.
LEPIDONOTINAE. Eighteen pairs of elytrae; 43 Notosetae few, thick, recurved uni- or bidentate with
segments. Lateral antennae subterminal. Notopodia coarse serrations. Neurosetae few, thick and falcate,
reduced with a few, nearly capillary setae. Neurosetae uni- or bidentate.
thin, slender, usually bifid.
Arctonoella Buzhinskaya 1967, Harmothoe sinaga-
Alentiana Hartman 1942, Polynoe aurantiaca Verrill waensis Izuka 1912; only species.
1885; only species. LEPIDONOTINAE. Sixteen pairs of elytrae, 41 seg-
LEPIDONOTINAE. Seventeen to 20 pairs of elytrae; ments. Lateral antennae subterminal. Notosetae thinner
36-39 segments. Notosetae absent. Neurosetae uni- than neurosetae, distally capillary. Neurosetae uni-
dentate, serrated or smooth. dentate.
Allmaniella McIntosh 1885, A. setubalensis McIntosh Australaugeneria Pettibone 1969d, Polynoe rutilans
1885; 5 species. Grube 1878; 2 species.
LEPIDONOTINAE. More than 15, but less than 30 HARMOTHOINAE. Fifteen pairs of elytrae, ap-
pairs of elytrae. Prostomium anteriorly produced into proximately 40 segments. Notosetae thicker than
two large lobes with the small lateral antennae attached neurosetae, smooth or faintly serrated. Neurosetae
distally. Notosetae thick and finely serrated; neuro- varied: as large golden hooks in the second and third
setae of two kinds, upper slender and smooth, lower segment; otherwise uni- or bidentate, often spurred
thicker, bidentate and vaguely serrated. and subdistally inflated.
Andresia Prenant 1924, A. ampullifera Prenant 1924; Austrolaenilla Bergstrom 1916, A. antarctica Berg-
only species. strom 1916; 6 species.
HARMOTHOINAE. Thirteen to 14 pairs of elytrae; HARMOTHOINAE. Fifteen to 16 pairs of elytrae;
32-33 segments. Notosetae thicker than neurosetae, 40-43 segments. Notosetae thicker than neurosetae
coarsely serrated. Neurosetae unidentate, serrated. with transverse rows of teeth. Neurosetae unidentate
Some notocirri with large, spherical ampullae sub- or bidentate, with the distal end pencillate.
distally.
Barrukia Bergstrom 1916, Gatryana cristata Willey
Antinoana Hartman and Fauchald 1971, A. fusca Hart- 1902; 2 species.
man and Fauchald 1971; only species. HARMOTHOINAE. Fifteen pairs of elytrae, 35-36
HARMOTHOINAE. Twelve to 13 pairs of elytrae; segments. Notosetae distally with tuft of long, fine
26-27 segments. Notosetae thicker than neurosetae, hairs, otherwise blunt and serrated. Neurosetae uni-
vaguely serrated. Neurosetae slender and bifid. dentate, with a few coarse teeth on the cutting edge.
Antinoe Kinberg 1855, A. microps Kinberg 1855; 8 Bathyadmetella Pettibone 1967, B. commando Petti-
species. bone 1967; only species.
HARMOTHOINAE. Fifteen pairs of elytrae. Noto- LEPIDONOTINAE. Twenty-three pairs of elytrae,
setae thicker than neurosetae, blunt, transversely ser- 58 segments. Notosetae absent, notopodia reduced.
rated. Neurosetae with slender tips, but not fdamentously Neuropodia prolonged, with slender, flattened setae,
prolonged. finely dentate and distally very abruptly tapering.
Antinoella Augener 1928b, Antinoe sarsi Kinberg in Bathymoorea Pettibone 1967, Polynoe ?renotuberculata
Malmgren 1865; 7 species. Moore 1910; only species.
HARMOTHOINAE. Fifteen pairs of elytrae; ap- LEPIDONOTINAE. Numbers of elytrae not known,
proximately 40 segments. Notosetae thicker than neuro- 35 segments. Notosetae short and slender, neurosetae
thicker, long, bidentate and marginally serrated. Renal HARMOTHOINAE. Fifteen pairs of elytrae; poste-
papillae greatly prolonged in some median setigers. rior half of body not covered by elytrae. Notosetae
mainly capillaries, a few thicker blunt setae present.
Benhamipolynoe Pettibone 1970d, Lepidasthenia anti-
pathicola Benham 1927; only species. Neurosetae thicker than notosetae, mainly unidentate,
but a few bidentates also present. Prostomium round,
LEPIDONOTINAE. More than 17 pairs of elytrae;
eyes conspicuous.
numerous segments. Notosetae absent. Neurosetae
thick, falcate with weakly marked serrations. Eucranta Malmgren 1865, E. villosa Malmgren 1865;
4 species.
Bouchiria Wesenberg-Lund 1949, B. vesiculosa
HARMOTHOINAE. Fifteen pairs of elytrae; 36-40
Wesenberg-Lund 1949; only species. segments. Notosetae thicker than neurosetae with rows
LEPIDONOTINAE. Numbers of pairs of elytrae of teeth. Some neurosetae distally split with both parts
and segments not known. Notosetae absent; neurosetae about equally long and thick; other neurosetae slender
include slender capillary setae with long slender spines and unidentate.
and thicker more acicular, but otherwise similar setae
ventrally. Palps reduced to two small tubercles. Nu- Eulagisca McIntosh 1885, E. corrientis McIntosh 1885;
merous stalked papillae present on the parapodia. 3 species.
LEPIDONOTINAE. Fifteen pairs of elytrae; 37 seg-
Bylgides Chamberlin 1919c, Bylgia elegans Theel ments. Lateral antennae subterminal; facial tubercle
1879; only species. present. Notosetae slender, numerous, pectinate, thicker
IPHIONINAE. Two antennae; facial tubercle absent. than the neurosetae. Neurosetae slender and tapering
Notosetae coarser than neurosetae. to capillary tips.
Cervilia Frickinger 1916, C. japonica Frickinger 1916; Eunoe Malmgren 1865, E. oerstedi Malmgren 1865;
only species. 40 species.
Sub-family unknown. Three antennae, lateral anten- HARMOTHOINAE. Fifteen pairs of elytrae, ap-
nae attached posteriorly on the prostomium. Fifteen proximately 40 segments. Notosetae thicker than neuro-
pairs of elytrae. Notosetae slender and capillary; neurosetae with rows of spines; neurosetae all unidentate,
setae unidentate and thicker. with more or less well-marked spinose region.
Chaetacanthus Seidler 1924, Iphione magnifica Grube Euphione McIntosh 1885, E. elisabethae McIntosh
1875; 3 species. 1885; 6 species.
LEPIDONOTINAE. Twelve pairs of elytrae, 26 seg- LEPIDONOTINAE. Twelve pairs of elytrae, short-
ments. Elytrophores with branchiae. Notosetae fine bodied. Elytrophores with branchiae. Notosetae cap-
and capillary; neurosetae unidentate and spinose. illary; neurosetae thicker, unidentate and laterally
covered with fine hairs.
Dilepidonotus Hartman 1967, D. falklandicus Hartman
1967; only species. Euphionella Monro 1936, Physalidonotus lobulatus
LEPIDONOTINAE. Twelve pairs of elytrae, 26 Seidler 1922; 3 species.
setigers. Notosetae slender and silky capillaries; neuro- LEPIDONOTINAE. Twelve pairs of elytrae; 25 seg-
setae thicker, smooth and pointed. Dorsum of cirral ments. Elytrophores with branchiae. Notosetae cap-
segments crested; pseudelytrae present. ill ary; neurosetae thicker, unidentate and smooth.
Pseudelytrae present.
Drieschia Michaelsen 1892, D. pelagica Michaelsen
1892; 6 species. Frennia Viguier 1912, F. dubia Viguier 1912; 2
LEPIDONOTINAE. Twelve to 13 pairs of elytrae; species.
26-27 segments. Notopodia absent; upper neurosetae LEPIDONOTINAE. Numbers of pairs of elytrae
capillary, lower thicker and slightly serrated. and segments not known. Notopodia completely re-
duced; neuropodia strongly prolonged with ventral
Drieschella Augener and Pettibone in Pettibone 19704, cirri attached near the middle. All setae slender, smooth
D. maculata Augener and Pettibone in Pettibone capillaries.
19704; only species.
LEPIDONOTINAE. Twenty pairs of small elytrae; Gastrolepidia Schmarda 1861, G. clavigera Schmarda
47 segments. Notosetae absent; neurosetae slender 1861; only species.
and tapering to capillary tips. Presetal lobes longer LEPIDONOTINAE. More than 21 pairs of elytrae.
than postsetal ones; acicular lobes not projecting. Notosetae and neurosetae similar in thickness; notosetae
blunt and serrated; neurosetae unidentate and serrated.
Enipo Malmgren 1865, E. kinbergi Malmgren 1865; Antennae and dorsal cirri strongly inflated subdistally,
4 species. with slender tips. Ventrum with large lamellae.

Gattyana McIntosh 1900, Aphrodita cirrhosa Pallas HARMOTHOINAE. Fifteen pairs of elytrae; poste-
1766; 14 species. rior part of body not covered by elytrae. Notosetae as
HARMOTHOINAE. Fifteen pairs of elytrae; ap- thick as neurosetae, vaguely serrated. Neurosetae dis-
proximately 40 segments. Most notosetae capillary with tally bi- or unidentate.
fine dentition. Neurosetae thicker than notosetae,
Herdmanella Darboux 1899, Polynoe ascidioides
distally unidentate, serrated.
McIntosh 1885; 3 species.
Gorekia Bergstrom 1916, Malmgrenia crassicirris HARMOTHOINAE. Eight to 9 pairs of elytrae;
Willey 1902; only species. 15-17 segments. Notosetae spinose, neurosetae less
HARMOTHOINAE. Fifteen pairs of elytrae; 38-40 so, both conspicuously long and unidentate, parapodia
segments. Notosetae shorter and thicker than the neuro- prolonged.
setae, finely serrated. Neurosetae in part distally trifid.
Hermadion Kinberg 1855, H. magalhaensi Kinberg
Grubeopolynoe Pettibone 1969b, Polynoe tuts Grube 1855; 5 species.
1855; only species. HARMOTHOINAE. Fifteen pairs of elytrae; more
HARMOTHOINAE. Fifty or more pairs of elytrae; than 50 segments. Notosetae thicker than neurosetae
numerous segments. Notosetae slenderer than neuro- and strongly serrated. Neurosetae uni- or bidentate,
setae; of two kinds, short and blunt and slender and serrated. Notosetae held erect over body.
tapering. Neurosetae all of one kind, vaguely bidentate
Hermenia Grube 1856, H. verruculosa Grube 1856;
or unidentate, serrated. Neuropodia with long supra-
2 species.
acicular postsetal lobe.
LEPIDONOTINAE. Twelve pairs of elytrae, small,
Halosydna Kinberg 1855, H. patagonica Kinberg 1855; not overlapping. Notosetae few, slender and serrated;
14 species. neurosetae distally trifurcate.
LEPiDONOTINAE. Eighteen pairs of elytrae; 37 seg-
Hesperonoe Chamberlin 1919, Harmothoe complanata
ments. Notosetae much finer than neurosetae, pointed
Johnson 1901; 4 species.
and serrated. Neurosetae thick, uni- or bidentate, with
HARMOTHOINAE. Fifteen pairs of elytrae; 36-38
coarse serrations.
segments. Notosetae in part at least as thick as neuro-
Halosydnella Hartman 1938, Halosydna australis setae; of two kinds; thick serrated and blunt and slender,
Kinberg 1855; 9 species. serrated and pointed. Neurosetae all unidentate, and
LEPH)ONOTINAE. Twenty-one pairs of elytrae; serrated, usually slender superior and thick in inferior
45 segments. Notosetae finer than neurosetae; serrated. positions.
Neurosetae distally uni- or bidentate, subdistally ser-
Heteropolynoe Bidenkap 1907, H. nordgaardi Biden-
rated.
kap 1907; only species.
Halosydnopsis Uschakov and Wit 1959, Halosydna HARMOTHOINAE. Numbers of pairs of elytrae
pilosa Horst 1917; only species. not known, 58 segments. Notosetae absent, all neuro-
LEPLDONOTINAE. Twenty-seven pairs of elytrae, setae unidentate and marginally serrated, slender in
body covered. Notosetae finer than neurosetae, finely superior and coarse in inferior positions.
serrated; neurosetae nearly smooth and unidentate.
Distal end of parapodia with series of large papillae. Hololepida Moore 1905, H. magna Moore 1905; 3
species.
Harmothoe Kinberg 1855, H. spinosa Kinberg 1855; LEPIDONOTINAE. Numerous pairs of elytrae,
120 species. numerous segments. Notosetae few in numbers, cap-
HARMOTHOINAE. Fifteen pairs of elytrae; ap- illary; neurosetae thicker, of two kinds, tanceotate
proximately 40 segments. Notosetae thicker than neuro- and bidentate with transverse rows of pockets. Nuchal
setae; with rows of spines. Neurosetae at least in part flap present.
bidentate, but usually also some unidentate in inferior
Hololepidella Willey 1905, H. commensalis Willey
positions.
1905; 5 species.
Hartmania Pettibone 1955, H. moorei Pettibone 1955; HARMOTHOINAE. Twenty-six or more pairs of
only species. elytrae; 55 or more segments. Notosetae at least as
HARMOTHOINAE. Fifteen pairs of elytrae; 40 seg- thick as neurosetae, nearly smooth with blunt tips.
ments. Notosetae finer than neurosetae, tapering to Neurosetae distally bidentate or entire.
sharp tips. Neurosetae tapering to sharp, pointed tips,
Hyperhalosydna Augener 1922, Lepidonotus striatus
not falcate.
Kinberg 1855; 2 species.
Hemilepidia Schmarda 1861, H. tuberculata Schmarda LEPIDONOTINAE. Twenty-one to 22 pairs of
1861; 4 species. elytrae; 50 segments. Notosetae few, short, curved

and blunt. Neurosetae bidentate with long, curved of accessory teeth in the crotch between the two major
tips. Lateral antennae terminal. teeth.
Intoshella Darboux 1899, Polynoe (Langerhansia) Lepidonotus Leach 1816, Aphrodita clava Montagu
euplectellae McIntosh 1885; 3 species. 1808; 65 species.
HARMOTHOINAE. Fifteen pairs of elytrae, ap- LEPIDONOTINAE. Twelve pairs of elytrae; 26 seg-
proximately 40 segments. Noto- and neurosetae similar ments. Notosetae finer than neurosetae, all tapering
in thickness; notosetae smooth-tipped, weakly ser- with whorls of spines. Neurosetae with rows of coarse
rated; neurosetae unidentate, more distinctly serrated. teeth; rarely bidentate, usually unidentate.
Eyes absent.
Leucia Malmgren 1867, Polynoe nivea Sars 1863;
Iphione Kinberg 1855, Polynoe muricata Savigny only species.
1818, 4 species. HARMOTHOINAE. Sixteen pairs of elytrae, short-
IPHIONINAE. Thirteen pairs of elytrae; notosetae bodied. Notosetae coarser than neurosetae and serrated.
capillary, neurosetae serrated and distally entire. Neurosetae long, slender, unidentate and serrated.
Kermadecella Darboux 1899, Polynoe magnipalpa Lucopia Pillai 1965, L. magnicirra Pillai 1965; only
McIntosh 1885; only species. species.
HARMOTHOINAE. Fifteen pairs of elytrae, short LEPIDONOTINAE. Fourteen pairs of elytrae; 27
body. Notosetae thicker than neurosetae, serrated. segments. Notosetae absent; neurosetae bidentate and
Neurosetae with transverse rows of spines; distally serrated. Dorsal cirri strongly inflated.
unidentate. Dorsal cirri alternating long and short, the
short ones basally inflated. Macellicephala McIntosh 1885, M. mirabilis McIntosh
1885; 18 species.
Lagisca Malmgren 1865, Polynoe rarispina Sars 1861; MACELLICEPHALINAE. Eight to 13 pairs of
24 species. elytrae; 17-29 segments. Notosetae few or absent, if
HARMOTHOINAE. Fifteen pairs of elytrae, ap- present then with marginal teeth. Neurosetae long,
proximately 50 segments. Noto- and neurosetae about usually paddle-shaped.
equally thick. Notosetae with dense rows of teeth;
neurosetae at least in part bidentate; all neurosetae Maceliicephaloides Uschakov 1955b, M. grandicirra
serrated. Posterior 8-10 segments not covered by Uschakov 1955b; only species.
elytrae. MACELLICEPHALINAE. Maximally 9 pairs of
elytrae; 16-17 segments. Notosetae absent. Neurosetae
Lepidasthenia Malmgren 1867, Polynoe elegans Gmbe long, marginally dentate. Prostomium with two very
1840; 37 species. strongly inflated lobes.
LEPIDONOTINAE. Numerous pairs of elytrae and
segments. Notosetae few, usually blunt; neurosetae Macelloides Uschakov 1957, M. antarctica Uschakov
numerous, uni- or bidentate, most are thicker than 1957; only species.
notosetae except inferior ones in each fascicle. Lateral MACELLICEPHALINAE. Fifteen pairs of elytrae;
antennae terminal. 30 segments. Notosetae absent. Neurosetae distally
inflated.
Lepidastheniella Monro 1924, Polynoe comma Thom-
son 1902; 3 species. Maimgrenia McIntosh 1874, M. whiteavesi McIntosh
LEPIDONOTINAE. Up to 90 pairs of elytrae; cover- 1874; 9 species.
ing the body. Notosetae thinner than neurosetae, ringed LEPIDONOTINAE. Fifteen pairs of elytrae; 36-41
with spines. Neurosetae spinose and distally entire. segments. Noto- and neuroseta similar in thickness.
Notosetae nearly smooth; neurosetae uni- or bidentate
Lepidofimbria Hartman 1967, L. oculata Hartman with very small secondary teeth. Lateral antennae
1967; only species. subterminal.
LEPIDONOTINAE. Numbers of pairs of elytrae
and segments not known. Ventrum with transverse Maimgreniella Hartman 1967, M. dicirra Hartman
row of three papillae on each segment; ventral citrus 1967; only species.
heavily funbriated. Notosetae assent; neuroseta smooth LEPIDONOTINAE. Fifteen pairs of elytra; 41-56
and distally entire. segments. Noto- and neuroseta similar in thickness;
notosetae falcate, nearly smooth. Neuroseta bidentate
Lepidogyra Hartman 1967, L. alba Hartman 1967;
with long, slender secondary tooth. Dorsal cirri of two
only species.
kinds; long and slender and short and expanded.
LEPIDONOTINAE. Numbers of pairs of elytrae
and segments not known. Notosetae coarser than neuro- Meiaenis Malmgren 1865, M. loveni Malmgren 1865;
seta; neuroseta spinose, distally bifid with a series only species.

HARMOTHOINAE. Fifteen pairs of elytrae; 39-41 enlarged dorsally; pre- and postsetal lobes of similar
segments. Notosetae thicker than neurosetae, few in length.
number; smooth or faintly structured. Neurosetae of
Phyllohartmania Pettibone 1961, P. taylori Pettibone
two kinds: numerous slender dentate with capillary
1961; only species.
tips; few furcates with subequal blunt tips.
HARMOTHOINAE. Fourteen pairs of elytrae, less
Nemidia Malmgren 1865, N. torelli Malmgren 1865; than 40 segments. Notosetae slender and spinose with
9 species. capillary tips; neurosetae similar in thickness; distally
HARMOTHOINAE. Fifteen pairs of elytrae; poste- spinose and spinigerous. Ventral surface with paired
rior part of body without elytrae. Notosetae mainly foliose appendages on each segment.
capillaries, but a few thick spines present; neurosetae Phyllosheila Pettibone 1961, P. wigleyi Pettibone 1961;
mainly unidentate, but a few bidentate setae present. only species.
Prostomium quadrangular, eyes missing or strongly HARMOTHOINAE. Fifteen pairs of elytrae; less
reduced. than 50 segments. Notosetae thicker than neurosetae,
spinose. Neurosetae smooth and distally bidentate.
Neohololepidella Pettibone 1969b, N. murrayi Petti-
Ventral cirri foliose, ventral surface papillated.
bone 1969b; only species.
HARMOTHOINAE. Fifty or more pairs of elytrae; Podarmus Chamberlin 1919c, P. ploa Chamberlin
numerous segments. Notosetae thicker than neurosetae, 1919c; 2 species.
nearly smooth, with blunt tips. Neurosetae with very LEPIDONOTINAE. Fourteen pairs of elytrae, 30
short bare tip beyond a dense spinose region; distally segments. Notosetae absent; neurosetae all distally
bidentate or entire. entire; of two kinds, thick and straight or slender and
capillary.
Paradyte Pettibone 1969a, Polynoe crinaidicola Potts
1910; 2 species. Polyeunoa McIntosh 1885, P. laevis McIntosh 1885;
HARMOTHOINAE. Fifteen pairs of elytrae; 40 seg- 3 species.
ments. Notosetae thicker than neurosetae, sabrelike HARMOTHOINAE. Nineteen to 30 pairs of elytrae,
with entire or slightly notched tips, nearly smooth. posterior part of body not covered by elytrae. Notosetae
Neurosetae of two kinds: supracicular ones with semi- thicker than neurosetae, faintly serrated. Neurosetae
lunar pockets, slender, spinose and with bifid tips. unidentate, subdistally expanded and dentate.
Subacicular ones thicker, with semilunar pockets and
Polynoe Savigny 1818, P. scolopendrina Savigny 1818;
entire tips.
17 species.
Parahalosydna Horst 1915a, P. sibogae Horst 1915a; HARMOTHOINAE. Fifteen pairs of elytrae; poste-
4 species. rior part of body not covered by elytrae. Notosetae
LEPIDONOTINAE. Fifteen pairs of elytrae; short- mainly thick and blunt-tipped, but a few capillary setae
bodied. Notosetae thinner than neurosetae, serrated. present. Nearly all neurosetae bidentate, except usually
Neurosetae unidentate and serrated along both edges. one or two unidentate in each of the posterior setigers;
most neurosetae coarser than the notosetae.
Parahololepidella Pettibone 1969b, Hololepidella
Polynoella McIntosh 1885, P. levisetosa McIntosh
greeffli Augener 1918; 2 species.
1 885; 3 species.
HARMOTHOINAE. Numerous pairs of elytrae and
HARMOTHOINAE. Twelve pairs of elytrae; 25-26
segments. Notosetae slenderer than neurosetae, but
segments. Notosetae absent. Neurosetae long, slender,
still thick. Neurosetae very thick, slightly hooked,
unidentate and falcate. Neuropodia long, pointed and
entire and very faintly spinose. Neuropodia with sub-
distally bifid.
acicular digitiform process.
Pottsiscalisetosus Pettibone 1969a, Scalisetosus prae-
Paralepidonotus Horst 1915a, Polynoe ampullifera longus Marenzeller 1902; only species.
Grube 1878; 4 species. HARMOTHOINAE. Twenty-eight or more pairs
LEPIDONOTINAE. Fifteen pairs of elytrae; 38 seg- of elytrae; numerous segments. Notosetae finer than
ments. Notosetae thicker than the neurosetae; densely neurosetae; tapered to blunt tips, serrated. Neurosetae
serrated. Neurosetae at least in part bidentate. with semilunar pockets, distally entire and more or
Perolepis Ehlers 1908, P. regularis Ehlers 1908; only less falcate.
species. Pseudohalosydna Fauvel 1913, P. rosea Fauvel 1913;
LEPIDONOTINAE. Numerous pairs of elytrae and only species.
segments. Notosetae absent; neurosetae distally bi- LEPIDONOTINAE. At least 20 pairs of elytrae;
dentate. Ventral cirri with three knobs; cirrophores numerous segments. Notosetae spinose and capillary;

neurosetae of two kinds; superior ones slender and bodied. Notosetae of two kinds; short and lancet-
spiralled spinose; inferior ones acicular and spinose. shaped, and longer, serrated and tapering. Neurosetae
serrated and unidentate.
Pseudopolynoe Day 1962, Polynoe inhacae Day 1951;
only species. Uncopolynoe Hartmann-Schroder 1960a, U. corallicola
LEPIDONOTINAE. Fifteen to 17 pairs of elytrae; Hartmann-Schrdder 1960a; only species.
posterior half of body not covered by elytrae. Notosetae HARMOTHOINAE. Numbers of elytrae not known,
finer than neurosetae, serrated; neurosetae bi- or uni- approximately 44 segments. Notopodia absent; first
dentate, serrated. three neuropodia with strongly curved hooks; others
with uni- or bidentate setae with serrated subdistal
Robertianella McIntosh 1885, R. synophthalma McIn-
areas.
tosh 1885; only species.
HARMOTHOINAE. Thirteen pairs of elytrae; ap- Weberia Horst 1915b, W. pustulate Horst 1915b; 3
proximately 30 segments. Noto- and neurosetae of species.
similar thickness; notosetae blunt, nearly smooth; neu- LEPIDONOTINAE. Eighteen pairs of elytrae; short-
rosetae distally minutely notched, nearly smooth. Eyes bodied. Notosetae absent; neurosetae unidentate and
very large and nearly confluent on the prostomium. curved. Ventral cirri absent except in second segment;
Scalisetosus McIntosh 1885, S. ceramensis McIntosh dorsum with a pustule on each segment.
1885; 8 species.
HARMOTHOINAE. Sixteen pairs of elytrae; 40
segments. Notosetae much coarser than neurosetae, Invalid Genera
with a few spines only. Neurosetae slender, distally
entire and spinose. Both noto- and neuropodia with Agnodice McIntosh 1885, see Lagisca
long lobes. Bathynoe Ditlevsen 1917, see Weberia
Bylgia Theel 1879, see Bylgides
Sheila Monro 1930, S. bathypelagica Monro 1930;
Chaetosphaera Haecker 1898, larvae of several genera
only species.
Dasylepis Malmgren 1867, see Acanthicolepis
LEPIDONOTINAE. Thirteen pairs of elytrae; 29
Eumolpe Oken 1807, see Lepidonotus
segments. Notosetae slender, dentate capillaries; neuro-
Eupolynoe McIntosh 1874, see Eucranta
setae of several kinds: numerous superior dentate cap-
Evarne, Malmgren 1865, see Harmothoe
ill aries; most of the others coarse, dentate and entire;
Evarnella Chamberlin 1919c, see Harmothoe
one very large bidentate hook in the middle of each
Gastroceratella Darboux 1899, see Thormora
fascicle.
Halosydnoides Seidler 1924, see Arctonoe
Subadyte Pettibone 1969a, Polynoe pellucida Ehlers Harmopsides Chamberlin 1919c, see Lepidasthenia
1864; 3 species. Iphionella McIntosh 1885, see Iphione
HARMOTHOINAE. Fifteen to 16 pairs of elytrae; Laenilla Malmgren 1865, see Harmothoe
approximately 40 segments. Notosetae similar in thick- Langerhansia McIntosh 1885, see Intoshella
ness to the neurosetae; with spinose pockets and slightly Lepidametria Webster 1879b, see Lepidasthenia
notched tips. Neurosetae with semilunar pockets, Nectochaeta Marenzeller 1892, in part Lepidasthenia,
spinose and distally bidentate. also generally polynoid larvae and juveniles
Norepa Baird 1865, see Iphione
Telolepidasthenia Augener and Pettibone in Pettibone
Nychia Malmgren 1865, see Gattyana
1970d, T. lobetobiensis Augener and Pettibone in
Pettibone 1970d; only species. Oligolepis Levinsen 1887, see Macellicephala
Paranychia Czerniavsky 1882, questionably Lagisca
LEPIDONOTINAE. More than 21 pairs of elytrae
Parapolynoe Czemiavsky 1882, see Polynoe
and more than 50 segments. Notosetae absent; neuro-
setae slender with spinose regions and entire tips. Parmensis Malmgren 1867, see Harmothoe
Physalidonotus Ehlers 1905, see Euphione
Presetal lobes longer than postsetal ones.
Plotolepis Chamberlin 1919c, see Drieschia
Tenonia Nichols 1969, T. kitsapensis Nichols 1969; Quetieria Viguier 1911, juvenile form
only species. Tricosmochaeta Morgera 1918, see Harmothoe
HARMOTHOINAE. Fifteen pairs of elytrae; ap-
proximately 40 segments. All setae similar in thick-
ness; all slender; notosetae capillary, neurosetae in
FAMILY POLYODONTIDAE BUCHANAN 1894
part bidentate.
Thormora Baird 1865, T. jukesii Baird 1865; 5 species. Aphroditaceans with dorsoventrally flattened bodies.
LEPBDONOTINAE. Twelve pairs of elytrae; short- Two or three antennae present, median antenna when

present, attached dorsally or posteriorly on the pros- The polyodontids are large, solid-bodied scale-
tomium (antennae may be absent). Eversible pharynx worms, usually covered with thick, loosely constructed
with four jaws. Marginally fringed or smooth elytrae tubes consisting of thin threads filled with clay or
alternate with dorsal cirri in all setigers. All setae sand particles. The most important recent revision
simple. Spinning glands present. was made by Strelzov (1968b).
Key to Genera
la. Ommatophores present 2
lb. Ommatophores absent 5
2a (Ia). Two antennae present 3
2b (Ia). Three antennae present 4
3a (2a). Ommatophores fused in the midline Neopanthalis
3b (2a). Ommatophores separated from each other Eupolyodontes
4a (2b). Superior neurosetae long and tapering, more or less hirsute Polyodontes
4b (2b). Superior neurosetae short and brush-topped Panthalis
5a (lb). Three antennae present Eupanthalis
5b (lb). Antennae absent Resno
Generic Definitions
Eupanthalis McIntosh 1876, E. kinbergi McIntosh Restio Moore 1903, R. aenus Moore 1903; only species.
1876; 8 species. Ommatophores and antennae absent. Setae include
Ommatophores absent, three antennae present; me- long, slender ones with awnlike spines, thick, colorless
dian antenna attached dorso-posteriorly. Setae include and tapering, slightly curved with pilose middle part,
slender, pilose tapering capillaries; bluntly acicular, acicular in appearance, thick, with broad lance-shaped
aristate ones with pilose shafts, and side-shaped, ends and transverse rows of fine hairs.
dentate ones.
Eupolyodontes Buchanan 1894, E. cornishii Buchanan Taxonomic Note

1894; 5 species.
Ommatophores present and separated from each Pseudeupanthalis Fauvel (1957), described in this
other. Two frontal antennae present; small nuchal family, appears to be synonymous with Sthenelanella
papilla present. Setae include bipinnate pencillate setae; in the family Sigalionidae.
serrulate setae and smooth, acicular spines.
Neopanthalis Strelzov 1968b, N. pelamida Strelzov Invalid Genera
1 968b; only species.
Ommatophores present but fused in the midline. Acoetes Audouin and Milne Edwards 1832, see
Lateral antennae short and inserted dorsally on the Polyodontes
ommatophores. Euarche Ehlers 1887, see Eupanthalis
Eupompe Kinberg 1855, see Polyodontes
Panthalis Kinberg 1855, P. oerstedii Kinberg 1855; Lepidia Savigny in Audouin and Milne Edwards 1832,
12 species. see Panthalis.
Ommatophores present and separated; three antennae
present. Setae include short brush-topped setae; thick,
blunt acicular setae with rows of teeth subdistally and Pholoididae NEw NAME
dentate side-shaped setae.
Aphroditaceans with flattened bodies. One median
Polyodontes Renier in Audouin and Milne Edwards antenna present. Four jaws present. Fringed elytrae
1832, Phyllodoce maxillosa Ranzani 1817; 12 alternate with dorsal cirri in all setigers. Each elytron
species. with concentric rings. Neurosetae composite, notosetae
Ommatophores present and separated; three antennae simple.
present. Setae include long, tapering hirsute ones; stout The family consists of one genus, Pholoides Pruvot
acicular setae with or without aristae, usually hirsute 1895 with genotype P. dorsipapillata Pruvot 1895.
or dentate; and dentate sickle-shaped setae. The family-name is to replace Peisidicidae Hartman
FIGURE 16. (A), Family PHOLOIDIDAE, Pholoides aspera, Mugu Submarine Canyon, California, about 100 m, 50x; (B),
elytron of the above, 50x; (C), Family EULEPETHIDAE, Grubeulepis fimbriata, Consag Rock, Golfo de California, elytron, l i x;
(D), diagram of the anterior end of the above, 8x; (E), median parapodium of the above, 25x; (F), anterior end of the above, first
two elytrophores removed, 10x.

and Fauchald 1971, based on the genus Peisidice John- FAMILY EULEPETHIDAE CHAMBERLIN 1 919c
son 1897. There appears to be no reason to maintain
Aphroditaceans with flattened bodies. Two antennae,
a separation between the two genera; the older name
four jaws present. Elytrae alternate with dorsal cirri
has priority, and the family name should be based on
in anterior setigers, present on all posterior setigers;
this generic name. Four species are presently con-
elytrae marginally notched or with flattened marginal
sidered valid.
lappets. All setae simple. Neuracicula distally hammer-
headed.
The eulepethids (formerly pareulepids SENsu Hart-
Invalid Genera
man) were recently the subject of a monographic study
Parapholoe Hartmann-Schroder 1965, see Pholoides by Pettibone (1969e). The present survey follows hers
Peisidice Johnson 1897, see Pholoides closely in most matters.
Key to Genera
Ia. Elytrae 12 pairs, increasing in length progressively, followed by posterior pseudelytrae 2
lb. Elytrae more than 12 pairs; the first 12 pairs increasing in length posteriorly; the more posterior pairs
smaller 3
2a (la). Elytrae with lateral border notched Pareulepis
2b (Ia). Elytrae with flattened lateral marginal lappets Grubeulepis
3a (I b). Elytrae with lateral borders notched Eulepethus
3b (lb). Elytrae with flattened lateral marginal lappets Mexieulepis
Taxonomic Note Pareulepis Darboux 1899, Eulepis wyvillei McIntosh

1855; 2 species.
The pseudelytrae referred to in the key are sessile
Body with about 37 segments. Twelve pairs of
in contrast to the true elytrae, which are equipped with
elytrae on segments 2,4,5,7 ... 21,24; all with lateral
distinct elytrophores. They are in the same general
border notched. Pseudelytrae begin on segment 28.
position as the elytrae and appear to have similar
functions.
Invalid Genus
Generic Definitions
Eulepis Grube 1875, see Eulepethus
Eulepethus Chamberlin 1919c, Eulepis hamifera Grube
1875; only species.
Body with 60-70 segments. Elytrae with marginal FAMILY SIGALIONIDAE MALMGREN 1867
notches on segments 2,4,5,7 ... 21,24 and from seg-
ment 28 small elytrae with entire margins on every Aphroditaceans with quadrangular or flattened bodies.
segment. One to three antennae present; four jaws present.
Marginally fringed elytrae alternate with dorsal cirri
Grubeulepis Pettibone 1969e, Eulepis funbriata Tread-
on anterior setigers and are present on all posterior
well 1901; 7 species.
setigers. Neurosetae composite, notosetae simple.
Body with about 38 segments. Elytrae with flattened
Sigalionids are common in soft bottoms; they tend
marginal lappets including in all twelve pairs, on
to be long-bodied and the scales are usually rather
segments 2,4,5,7 . . . 21,24; pseudelytrae begin on
closely appended to the bodies. This, combined with
segments 26-29.
the well-developed notopodial fascicles of setae, tend
Mexieulepis Rioja 1961, M. elongata Rioja 1961; to give them a rather quadrangular appearance. The
2 species. sigalionids are presently under revision by Pettibone,
Body with about 50 segments. Elytrae with flattened who has already published a series of important mono-
marginal lappets, including numerous pairs, on seg- graphs (Pettibone 1969c, 1970a, 1970b, 1970e, 1971b,
ments 2,4,5,7 ... 21,24; smaller elytrae on every 1971c). The key given below must be considered
segment starting on segments 27 or 28. provisional.
Key to Genera
Ia. One median antenna present 2

lb. At least two antennae present 3

FIGURE 17. Family SIGALIONIDAE, Sthenolepis japonica, Bay of Nha Trang, Viet Nam, 50 m, 25x.
2a (I a). Composite neurosetae with long, slender, articulated appendages Mayella

2b (la). Composite neurosetae short, smooth unidentate falcigers Pholoe
3a (1b). Two antennae present Sigalion
3b (lb). Three antennae present 4
4a (3b). Lateral antennae on the prostomium proper; all antennae with short ceratophores or short and incon-
spicuous 5
4b (3b). Lateral antennae fused to first setiger; median antenna with large median ceratophore 6
5a (4a). Eyes large, antennae with ceratophores Euthalenessa
5b (4a). Eyes small, antennae without ceratophores Thalenessa
6a (4b). Median ceratophore without auricles 7
6b (4b). Median ceratophore with auricles 10
7a (6a). Dorsal cirri absent on setiger 3; elytrae not sand-incrusted 8
7b (6a). Dorsal cirri present on setiger 3; elytrae sand-incrusted 9
8a (7a). Neurosetae composite spinigers with short, canaliculated appendages Leanira
8b (7a). Neurosetae composite, unidentate falcigers with marginally serrated appendages Pareupholoe
9a (7b). Elytrae small, leaving the middle of the dorsum uncovered; neurosetae with short bidentate ap-
pendages Euphofoe
9b (7b). Elytrae covering the dorsum, neurosetae with uni- or bidentate appendages of varying lengths
. . . .... .. . .. .. .. . . .. Psammolyce
IOa (6b). Auricles
. . . . .large,
. . . . .auricles
. . . . . .also
. . . present
.... on the tentacular . . . segment
. . . . . . . . . . . . . . . . . . . . . . . . . .Horstileanira
l0b (6b). Auricles small, absent from tentacular segment 11
l la (lob). All neurosetae composite unidentate falcigers with relatively short, straight appendages . . Sthenelanella
IIb (lob). At least some neurosetae otherwise (bidentate, or with articulated, canaliculated or spinigerous
appendages) 12
12a (1 Ib). Ventral cirri covered medially by long papillae Willeysthenelais
12b (I lb). Ventral cirri smooth 13
13a (126). Parapodial lobes and stylodes covered with fine fimbriae Fimbriosthenelais
13b (12b). Parapodial lobes and stylodes smooth 14
1 4a (13b). Long dorsal cirri on setiger 3 Neoleanira
14b (13b). Dorsal cirri absent from setiger 3 15

15a (14b). At least some neuropodial falcigers present Sthenelais

15b (14b). All neurosetae spinigerous 16
16a (15b). Dorsal tubercle present on setiger 3 Sthenolepis
lob (156). Dorsal tubercle absent from setiger 3 Ehlersileanira
Generic Definitions Pareupholoe Hartmann-Schroder 1962a, P. fimbriatus

Ehlersileanira Pettibone 1970c, Sthenelais incisa Grube Hartmann-Schroder 1962a; only species.
1877; 2 species. Three antennae; auricles absent. No dorsal cirri on
Three antennae; auricles present. Dorsal cirrus on setiger 3. Neurosetae composite, unidentate falcigers
setiger 3 absent. All neurosetae spinigerous, with with serrated cutting edge.
canaliculated appendages. Pholoe Johnston 1839, Aphrodita minuta Fabricius
Eupholoe McIntosh 1885, E. philippinensis McIntosh 1780; 10 species.
1885; only species. One antenna; auricles absent. Neurosetae composite
Three antennae; auricles absent. Dorsal cirrus on falcigers with short, unidentate appendages. Notosetae
setiger 3 present. Elytrae small, leaving middle of in part geniculate and strongly dentate.
body uncovered. Neurosetae with short, bidentate
Psammolyce Kinberg 1855, P. flava Kinberg 1855;
appendages. Encrusted with sand.
26 species.
Euthalenessa Darboux 1899, Thalenessa digitata McIn- Three antennae, auricles absent. Third setiger with
tosh 1885; 8 species. long dorsal cirri. Neurosetae composite falcigers with
Three small antennae, all on the prostomium, cerato- uni- or bidentate appendages of varying lengths.
phores present. Two pairs of large eyes. Neuropodia Elytrae cover body, sand-incrusted.
with large foliose lobes anteriorly. Neurosetae slender,
Sigalion Audouin and Milne Edwards 1832, S.
composite with multiarticulated distally bidentate
mathildae Audouin and Milne Edwards 1832; 12
appendages.
species.
Fimbriosthenelais Pettibone 1971b, Sthenelais longi- Two lateral antennae; auricles absent. Neurosetae
pinnis Grube 1870a; 2 species. simple and spinose or composite with appendages of
Three antennae; auricles present. No dorsal cirrus varying lengths, multiarticulated and bidentate.
on setiger 3. All parapodial lobes and stylodes
fimbriated. Neurosetae either simple and spinose or Sthenelais Kinberg 1855, S. helenae Kinberg 1855;
composite with distally bidentate, short, slender and 45 species.
articulated appendages. Three antennae, auricles present. No dorsal cirrus
on setiger 3'. Neurosetae include simple spinose and
Horstileanira Pettibone 1970a, H. vanderspoeli Petti- two kinds of falcigers, stout with short, bidentate
bone 1970a; 2 species. appendages, and slenderer falcigers with articulated
Three antennae, large auricles, auricles present also appendages.
on the tentacular segment. Setiger 3 with prominent
dorsal tubercle. Neurosetae simple and spinose and Sthenelanella Moore 1910, S. uniformis Moore 1910;
composite spinigers with long, canaliculated appen- 4 species.
dages; a few may be bidentate. Three antennae; auricles present. No dorsal cirri on
setiger 3. All neurosetae with short, unidentate ap-
Leanira Kinberg 1855, L. quatrefagesi Kinberg 1855; pendages.
11 species.
Three small antennae; auricles absent. No dorsal Sthenolepis Willey 1905, Leanira japonica McIntosh
cirrus on setiger 3. Neurosetae composite spinigers 1885; 20 species.
with relatively short, canaliculated appendages. Three antennae; auricles present. With dorsal tu-
bercles on setiger 3. Neurosetae all composite spinigers
Mayella Hartmann-Schroder 1959, M. articulata with canaliculated appendages.
Hartmann-Schroder 1959; only species.
One antenna, auricles absent. Neurosetae composite Thalenessa Baird 1868, Sigalion edwardsi Kinberg
spinigers with articulated appendages. 1855; 8 species.
Three small antennae, all on the prostomium, cerato-
Neoleanira Pettibone 1970a, Sigalion tetragonum phores absent. Two pairs of small eyes. Neurosetae
Orsted 1845; 2 species.
composite falcigers and a few simple spinose setae.
Three antennae; auricles present. Long dorsal cirri
on setiger 3. Neurosetae composite spinigers with Willeysthenelais Pettibone 1971b, Sthenelais diplocir-
canaliculated, relatively short appendages. rus Grube 1875; only species.

Three antennae; auricles present. No dorsal cirri Polylepis Grube 1878, see Psammolyce
on setiger 3. Neurosetae simple spinose or composite Pseudeupanthalis Fauvel 1957, see Sthenelanella
falcigers with thick, short appendages or more slender
with articulated appendages. Ventral cirri covered Superfamily Chrysopetalacea
medially with long papillae. Elytrae absent; notosetae flattened and expanded
paleae covering the dorsum. Prostomium not fused
Invalid Genera to the first setiger.
Conconia Schmarda 1861, see Sthenelais FAMILY CHRYSOPETALIDAE EHLERS 1864

Euleanira Horst 1916a, see Sthenelanella Chrysopetalaceans with short or long bodies, usually
Eusigalion Augener 1918, see Thalenessa strongly flattened. Three antennae. Notosetae in trans-
Eusthenelais McIntosh 1876, see Sthenelais and verse rows, held erect over the dorsum or covering
Leanira the back as tiles on a roof. Neurosetae composite.
Haswellia Darboux 1899, see Euthalenessa Chrysopetalids and palmyrids are very similar; the
Lepidopleurus Claparede 1868, see Psammolyce two families are often considered synonymous, in
Pelogenia Schmarda 1861, see Psammolyce which case the name Palmyridae applies (cf. Day 1967).
E
FIGURE 18. (A), Family CHRYSOPETALIDAE, Chrysopetalum occidentale, off Santa Catalina Island, California, 50 m, whole
animal, lOx; (B), anterior end of the above, setae of four first notopodia removed on the right side, 50x; (C), notoseta (palea) of
the above, 200x; (D), Family PALMYRIDAE, Palmyra aurifera, Eniwetok, Marshall Islands, shallow subtidal, notosetae removed
on four first segments on the right side and on the first segment on the left, 75x; (E), notoseta (palea) of the above, 50x.

Key to Genera
Ia. Notosetae cylindrical rather than flattened, not covering dorsum Dysponetus
lb. Notosetae flattened paleae 2
2a (lb). Prostomium without a conspicuous caruncle, body long, consisting of many segments ... Bhawania
2b (lb). Prostomium with a conspicuous caruncle overlapping the peristomium; body short 3
3a (2b). First segment with asymmetrical ventral cirri; dorsal palette of two abruptly different kinds
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . ............................................. Paleanotus
3b (2b). First segment with paired similar cirri; dorsal paleae of one kind only Chrysopetalum
Generic Definitions Superfamily Pisionacea
Bhawania Schmarda 1861, B. myriolepis Schmarda Elytrae absent; parapodia sub-biramous or uni-
1 861; 9 species. ramous; prostomium deeply imbedded in the first
Body with up to 300 segments, completely covered segment, or projecting freely between the first
by paleae. Caruncle absent, prostomium retractile segments.
under a fold from the first setigers. Palette of one or
t wo kinds, broad and narrow; neurosetae composite FAMILY PISIONIDAE SOUTHERN 1914
falcigers with blades of varying lengths.
Pisionaceans with maximally two pairs of antennae
Chrysopetalum Ehlers 1864, Palmyra debilis Grube on the prostomium. First segment with two pairs of
1 855; 5 species. tentacular cirri. Four jaws present. Setae composite
Body with about 40 segments, completely covered and simple; dorsal and ventral cirri usually clavate.
by paleae. Caruncle present. Paleae of one kind only; Siewing (1953), Laubier (1967b) and Stecher (1968)
first segment with paired, similar ventral cirri. recently have reviewed the family in terms of the
generic sub-division. Pisionura Hartman and Fauchald
Dysponetus Levinsen 1879, D. pygmaeus Levinsen
(1971) does not belong to this family (Hartmann-
1879; 4 species.
Schriider 1975). The key below is after Laubier (1967b).
Body with few segments, not covered by palette.
Caruncle absent. Notosetae cylindrical and erect over
the dorsum.
Paleanotus Schmarda 1861, P. chrysolepis Schmarda
1861; 6 species.
Body with about 40 segments, completely covered
by paleae. Caruncle may be present. Paleae of two
kinds, abruptly differing in shape. First segment with
strongly asymmetrical ventral cirri.
Invalid Genera
Heteropale Johnson 1897, see Paleanotus
Psectra Grube 1868a, see Bhawania
Taphus Webster and Benedict 1887, see Dysponetus
FAMILY PALMYRIDAE KINBERG 1858
Chrysopetalaceans with short, flattened bodies. One

antenna present. Notosetae in rosettes on the notopodial
tubercles. Neurosetae composite.
The palmyrids are known for one genus, Palmyra
Savigny 1818 with genotype P. aurifera Savigny 1818
and possibly for one or two more species; most de-
scribed forms are considered synonyms of P. aurifera
or indeterminable. The family is limited to shallow
water, usually sandy or generally hard substrates in
warm water and appear to be most common in the Room 19. Family PISIONIDAE, Pisione oerstedi, Indepen-
western Pacific Ocean. dencia Bay, Peru, shallow water, 25x.

Key to Genera
Ia. Median unpaired antennae present pisionella
l b. Median unpaired antennae absent 2
2a (1 b). With two pairs of similar cephalic appendages Pisionidens
2b (lb). With three pa irs of cephalic appendages of different structure 3
3a (2b). First segment asetigerous and apodous; proboscis unarmed Anoplopisione
3b (2b). No apodous and asetigerous segment present; proboscis with four jaws Pisione
Generic Definitions Suborder Nereidiformia

Anoplopisione Laubier 1967b, A. minuta Laubier
Phyllodocida with at least one pair of antennae; at
1967b; only species. least one pa ir of tentacular cirri; palps short and usu-
Pisionids with palps and two pairs of tentacular cirri ally distally blunt, frequently biarticulated. Eversible
present. Proboscis unarmed; first segment without pharynx, if armed, with one pair of lateral jaws and
parapodia and setae. sometimes with accessory denticles. First parapodia
Pisione Grube 1857, P. oerstedii Grube 1857; 11 lateral.
species.
Pisionids with palps and two pairs of tentacular cirri
present. Proboscis with four jaws; all anterior segments FAMILY HESIONIDAE SARS 1862
with parapodia and setae.
Relatively short-bodied, dorsoventrally flattened
Pisionella Hartman 1939, P. hancocki Hartman 1939; worms. Two or three antennae (antennae rarely absent);
only species. palps may be absent or have from one to three articles.
Pisionids with palps; two pairs of tentacular cirri Two to eight pairs of tentacular cirri present. Jaws may
and a median unpaired antenna present. Proboscis be present. Parapodia uniramous or biramous, but noto-
with four jaws; all anterior segments with parapodia podia always reduced compared to the neuropodia.
and setae. Dorsal cirri long and slender. Neurosetae composite;
Pisionidens Aiyar and Alikunhi 1943, Pisionella indica
notosetae, if present, simple.
The hesionids are one of the least known families
Aiyar and Alikunhi 1940; only species.
Pisionids with two pairs of similar cephalic appen- of polychaetes; the generic sub-division suggested by
the key below is very tentative; a large number of
dages. Proboscis with four jaws; all segments with
parapodia and setae. additional genera may be expected described within a
few years. Hesionids are common animals in hard
Invalid Genera
substrates and in shallow water; they are more rarely
found in deep water. They tend to be fragile and frag-
Fauveliella Tebble 1953, see Pisionidens ment easily upon collection; generally hesionids should
Pisionella Aiyar and Alikunhi 1940, see Pisionidens be handled as scale-worms: each specimen preserved
Praegeria Southern 1914, see Pisione separately.
Key to Genera
la. Two pairs of tentacular cirri; five antennae Hesiosyllis
lb. At least three pairs of tentacular cirri; maximally three antennae 2
2a (Ib). Four or more pairs of tentacular cirri 3
2b (lb). Three pairs of tentacular cirri 7
3a (2a). Five or more pairs of tentacular cirri 4
3b (2a). Four pairs of tentacular cirri 8
4a (3a). Five pairs of tentacular cirri Friedericiella
4b (3a). Six or more pairs of tentacular cirri 5
5a (4b). Seven or more pairs of tentacular cirri 6
5b (4b). Six pairs of tentacular cirri 12
6a (5a). Seven pairs of tentacular cirri Periboea
6b (5a). Eight pairs of tentacular cirri 22
7a (2b). Parapodia uniramous Orseis
7b (2b). Parapodia biramous Alikunhia

8a (3b). Parapodia distinctly uniramous 9

8b (3b). Parapodia sub-biramous or biramous 10
9a (8a). First setiger with large hooks Struwela
9b (8a). First setiger without hooks Hesiocaeca
Ills (8b). Antennae absent Bonuania
10b (8b). Antennae present 11
I la (l0b). Tentacular cirri on three segments (1-2-1); dorsal cirri articulated Hesionella
Frouae 20. (A), Family HESIONIDAE, Hesione intertexta, Puerto Rico, intertidal, 10x; (B), Family SYLLIDAE, Typosyllis
armillaris, El Descanso, Baja California, intertidal, median parapodium, 50x; (C). anterior end of the above, 50x; (D), Family
PILARGIIDAE, Sigambra bassi, outer harbor, Los Angeles, California, 50 m, 15x; (E), median parapodium of the above, 50x;
(F), Family CALAMYZIDAE, ? Calamyzas sp., diagrammatic outline from the dorsal side, 50x.

I lb (lob). Tentacular cirri on two segments (2-2); dorsal cirri smooth Hesionides
12a (5b). All parapodia uniramous 13
12b (5b). At least some parapodia sub-biramous or biramous 17
13a (12a). First three segments dorsally reduced Syllidia
13b (12a). Maximally first segment dorsally reduced 14
14a (13b). Tentacular cirri on four segments (first reduced) so that they appear as 3-2-1 Syllidia"
14b (13b). Tentacular cirri on three segments (2-2-2) 15
l5a (14b). Pharynx distally with a circlet of fine fimbriae Parasyllidea
15b (14b). Pharynx with either eleven or twenty-one distal papillae 16
16a (15b). Pharynx with 21 distal papillae Neopodarke
l6b (15b). Pharynx with eleven distal papillae Micropodarke
17a (12b). Three antennae present 18
l7b (12b). Two antennae present 21
18a (17a). Median antenna attached medially or posteriorly on the prostomium Microphthalmus
18b (17a). Median antenna attached frontally 19
19a(18b). Palpi simple Heteropodarke
19b (18b). Palpi biarticulated 20
20a (19b). Setae present from the second segment Ophiodromus
20b (19b). Setae present from the fourth segment Podarke"
21a (17b). Dorsal cirri smooth Parahesione
21b (17b). Dorsal cirri articulated Nereimyra
22a (6b). Three antennae 23
22b (6b). Two antennae 27
23a (22a). Median antenna attached medially or posteriorly on the prostomium 24
23b (22a). Median antenna attached frontally 26
24a (23b). Eversible pharynx distally fimbriated Amphiduros
24b (23b). Eversible pharynx distally papillated 25
25a (24b). Parapodia uniramous Leocratides
25b (24b). Parapodia biramous Leocrates
26a (23b). Setae present from the second segment Gyptis
26b (23b). Setae present from the fifth (or fourth) segment Podarkeopsis
27a (22b). Notopodia with falcate spines Hesiospina
27b (22b). Parapodia uniramous 28
28a (27b). Palps absent Hesione
28b (27b). Palps present 29
29a (28b). Setae present from the fourth segment Wesenbergia
29b (28b). Setae present from the third segment 30
30a (29b). Eversible pharynx smooth Dalhousiella
30b (29b). Eversible pharynx distally fimbriated Kefersteinia
Generic Definitions Bonuania Pillai 1965, B. parva, Pillai 1965; only

species.
Alikunhia Hartman 1959, Anophthalmus erythraeus
Antennae absent; biarticulated palps and four pairs
Alil unhi 1949; 4 species.
of tentacular cirri present. Parapodia biramous.
Three antennae, palps and three pairs of tentacular
cirri present. Parapodia biramous. Eversible pharynx Dalhousiella McIntosh 1901, D. carpenteri McIntosh
distally papillated, jaws absent. 1901; 5 species.
Two antennae; biarticulated palps and eight pairs
Amphiduros Hartman 1958, Amphidromus setosus
of tentacular cirri present. Parapodia uniramous, first
Hessle 1925; 3 species.
setae in third segment. Eversible pharynx distally
Three antennae, biarticulated palps and eight pairs
smooth, jaws absent.
of tentacular cirri present (2-2-2-2). Parapodia bi-
ramous. Eversible pharynx distally fimbriated, jaws Friedericiella Laubier 1967c, Hesionella pacifica
absent. Median antenna attached medially. Friedrich 1956; only species.

Three antennae, simple palps and five pairs of Two antennae, biarticulated palps and eight pairs
tentacular cirri present. Parapodia biramous, first of tentacular cirri present. Parapodia uniramous, setae
setae in fourth segment. from fourth segment. Eversible pharynx with distal
circlet of fimbriae; jaws absent.
Gyptis Marion and Bobretzky 1875, G. propinqua
Marion and Bobretzky 1875; 16 species. Leocrates Kinberg 1866b, L. chinensis Kinberg 1866b;
Three antennae, biarticulated palps and eight pairs 11 species.
of tentacular cirri present. Parapodia biramous; first Three antennae, biarticulated palps and eight pairs
setae in second segment. Eversible pharynx with 40 of tentacular cirri present. Parapodia biramous. Ever-
distal papillae, jaws absent. sible pharynx with jaws. Median antenna attached
posteriorly.
Hesiocaeca Hartman 1965, H. bermudensis Hartman
1965; only species. Leocratides Ehlers 1908, L. filamentosa Ehlers 1908;
Three antennae, biarticulated palps and four pairs only species.
of tentacular cirri present. Parapodia uniramous, first Three antennae, biarticulated palps and eight pairs
setae in third segment. Eversible pharynx with a few of tentacular cirri present. Parapodia uniramous. Ever-
distal papillae, jaws absent. sible pharynx with jaws. Median antenna attached
Hesione Savigny 1818, H. splendida Savigny 1818;
posteriorly.
7 species. Microphthalmus Mecznikow 1865, M. sczelkowii
Two antennae, palps absent, eight pairs of tentacular Mecznikow 1865; 13 species.
cirri present. Parapodia uniramous, setae first in third Three antennae, simple palps and six pairs of ten-
segment. Eversible pharynx distally smooth, jaws tacular cirri present. Parapodia sub-biramous. Eversible
absent. pharynx with distal circlet of papillae, jaws absent.
Hesionella Hartman 1939b, H. mccullochae Hartman Median antenna attached posteriorly.
1939b; only species. Micropodarke Okuda 1938, Kefersteinia dubia Hessle
Two antennae, palps absent, four pairs of tentacular 1925; only species.
cirri present. Parapodia sub-b iramous. Two antennae, biarticulated palps and six pairs of
Hesionides Friedrich 1937, H. arenaria Friedrich tentacular cirri present. Parapodia unramous. Eversible
1937; 3 species. pharynx with 11 papillae distally, jaws absent.
Three antennae, biarticulated palps and four pairs Neopodarke Hartman 1965, N. woodsholea Hartman
of tentacular cirri present. Parapodia sub-biramous to 1965; only species.
biramous, first setae in third segment. Eversible pharynx Two antennae, biarticulated palps and six pairs of
with ten distal papillae and two longer cirri, jaws tentacular cirri present. Parapodia uniramous; first
absent. setae in fourth segment. Eversible pharynx with 21
Hesiospina Imajima and Hartman 1964, Kefersteinia distal papillae, jaws absent.
similis Hessle 1925; only species.
Nereimyra Blainville 1828, Nereis punctata O.F.
Two antennae, biarticulated palps and eight pairs Muller 1776; 12 species.
of tentacular cirri present. Parapodia biramous. Ever- Two antennae, biarticulated palps and six pairs of
sible pharynx with 21-27 distal papillae; jaws absent. tentacular cirri present. Parapodia sub-biramous, first
Notopodial falcate spines present. setae in fourth segment. Eversible pharynx with circlet
Hesiosyllis Wesenberg-Lund 1950, H. enigmatica of papillae distally, jaws present.
Wesenberg-Lund 1950; only species.
Ophiodromus Sars 1862, Nereis fexuosa delle Chiaje
Five antennae (four frontal, one dorsal), smooth
1825; 11 species.
palps and two pairs of tentacular cirri present. Parapodia
Three antennae, biarticulated palps and six pairs of
biramous. Eversible pharynx with ten distal papillae;
tentacular cirri present. Parapodia biramous, first
jaws and teeth present.
setae in second segment. Eversible pharynx distally
Heteropodarke Hartmann-Schroder 1962a, H. hetero- with many fine frmbriae; jaws absent.
morpha Hartmann-Schroder 1962a; only species.
Orseis Ehlers 1864, O. pulls Ehlers 1864; 5 species.
Three antennae, smooth palps and six pairs of ten-
tacular cirri present. Parapodia sub-b iramous. Three antennae, simple palps and three pairs of
tentacular cirri present. Parapodia uniramous, first
Kefersteinia Quatrefages 1865, Psamathe cirrata setae in second segment. Median antenna attached
Keferstein 1862; only species. posteriorly.

Parahesione Pettibone 1956, Podarke luteola Webster Taxonomic Notes

1880; 2 species.
The genus Podarke usually is considered a synonym
Two antennae, simple palps and six pairs of ten-
of Ophiodromus. However, the concept as usually
tacular cirri present. Parapodia biramous, first setae
used, differs slightly from Ophiodromus, and has been
in second segment. Eversible pharynx with distal
included in the key and in the definitions. The same
circlet of fimbriae, jaws absent.
is true of the concept Syllidia as it has been used in the
Parasyllidea Pettibone 1961, P. humesi Pettibone literature, compared to the version originally proposed.
1 961; only species. The key is wholly dependent on correct identification
Two antennae, biarticulated palps and six pairs of of the numbers of pairs of tentacular cirri. This is per-
tentacular cirri present. Parapodia uniramous; first haps best observed on the lateral side of the animal,
setae in third segment. Eversible pharynx with a distal and with the light coming in under a low angle, so that
circlet of fimbriae, jaws absent. First segment reduced. each tentacular citrus will cast a shadow. Most hesionids
have dehiscent antennae and tentacular cirri so this
Periboea Ehlers 1864, P. tongocirrata Ehlers 1864;
may be the only means of getting the numbers of ante-
only species.
rior appendages correctly identified.
Two antennae, triarticulated long palps and seven
Hesionids in general are rather fragile, and have to
pairs of tentacular cirri present. Parapodia sub-biramous,
be handled very carefully in collections; the only
first setae in third segment. Eversible pharynx with
exceptions are the l arge, and rather solid-bodied mem-
1 6-22 distal papillae, jaws absent.
bers of the genus Hesione.
Podarke Ex AUCTORE; confused, possibly about 12
species.
Invalid Genera
Three antennae, biarticulated palps and six pairs of
tentacular cirri present. Parapodia biramous, first setae Anophthalmus Alikunhi 1949, see Alikunhia
from fourth segment. Eversible pharynx with a circlet Anoplonereis Giard 1882, see Ophiodromus
of papillae, ? jaws absent. Castalia Savigny 1820, see Nereimyra
Cirrosyllis Schmarda 1861, indeterminable
Podarkeopsis Laubier 1961b, P. galangaui Laubier Dalhousia McIntosh 1885, see Leocrates
1961b; only species. Hallmede Rathke 1843, see Nereimyra
Three antennae, biarticulated palps and eight pairs Mania Quatrefages 1865, see Ophiodromus
of tentacular cirri present. Parapodia biramous, first Orthodromus Ehlers 1908, see Ophiodromus
setae on fourth visible segment (fifth segment). Ever-
Oxydromus Grube 1855, see Gyptis
sible pharynx with circlet of papillae, jaws absent.
Pseudosyllidia Czerniavsky 1882, unidentifiable
Struwela Hartmann-Schroder 1959, S. noodti Hamnann- Schmardiella Czemiavsky 1882, unidentifiable
Schroder 1959; only species. Stephania Claparede 1870b, see Ophiodromus
Two antennae, biarticulated palps, four pairs of Telamone Claparede 1868, see Hesione
tentacular cirri present. Uniramous parapodia, first Tyrrhena Claparede 1868, see Leocrates
setae in third segment. First setiger with large hooks.
Syllidia Quatrefages 1865, S. armata Quatrefages FAMILY PILARGIIDAE SAINT-JOSEPH 1899

1865; 5 species. Nereidiforms with ribbon-shaped or cylindrical
Two antennae, biarticulated palps and six pairs of bodies. Two or three antennae present (rarely absent).
tentacular cirri present. Parapodia uniramous, first Palps simple or biarticulated; two pairs of tentacular
setae in second segment. Eversible pharynx with jaws. cirri (rarely absent). Proboscis unarmed. Parapodia
First to third segment dorsally reduced. biramous, but notopodia always reduced. Setae simple;
Syllidia ExAucroRe; confused, possibly five species. notosetae sometimes as a thick spine or hook. Notosetae
Two pairs of antennae, biarticulated palps and six may be absent.
pairs of tentacular cirri present. Parapodia uniramous, Pilargiids have been the subject of two recent revi-
first setae in fourth segment. sions on the generic level, Pearson (1970), and Emerson
and Fauchald (1971). Pettibone (1966b) added to the
Wesenbergia Hartman 1955, Hesionella problematica generic descriptions and revised several species. Pi-
Wesenberg-Lund 1950; only species. l argiids are never really numerous, but are present
Two antennae, biarticulated palps and eight pairs in most areas in moderate numbers; they tend to be
of tentacular cirri present. Parapodia uniramous, first associated with shelf depths and moderately coarse
setae in fourth segment. to mixed sediments.

Key to Genera
la. Notopodia with stout emergent hooks or spines 2

lb. Notopodia without emergent hooks or spines 8
2a (la). Notopodia with recurved emergent hooks 3
2b (Ia). Notopodia with stout, straight spines 6
3a (2a). Peristomium dorsally entire Ancistargis
3b (2a). Peristomium dorsally incised 4
4a (3b). Dorsal and ventral cirri reduced or absent; parapodia reduced, body subcylindrical Cabira
4b (3b). Dorsal and ventral cirri well developed, parapodia well developed, body dorso-ventrally flattened . 5
5a (4b). Antennae shorter than palps; integument papillated Ancistrosyllis
5b (4b). Antennae longer than palps, integument smooth Sigambra
6a (2b). Prostomial antennae present; parapodia sharply set off from body Synelmis
6b (2b). Prostomial antennae absent; parapodia distinct, but not set off from body 7
7a (6b). Tentacular cirri present; palps absent Litocorsa
7b (6b). Tentacular cirri absent, palps present Parandalia
8a (Ib). Prostomial antennae and tentacular cirri absent Loandalia
8b (lb). Prostomial antennae and tentacular cirri present 9
9a (8b). Prostomium with two antennae, biarticulate palps present Pilargis
9b (8b). Prostomium with three antennae; palps without palpostyles Otopsis
Generic Definitions Body flattened; three antennae and two pairs of

tentacular cirri present; palps present, but not articu-
Ancistargis Jones 1961, A. papillosus Jones 1961; 4
lated. Emergent spines absent.
species.
Body flattened, with two antennae, biarticulate palps, Parandalia Emerson and Fauchald 1971, P. ocularis
two pairs of tentacular cirri. Peristomium dorsally en- Emerson and Fauchald 1971; 6 species.
tire. Antennae shorter than palps. Emergent notopodial Body cylindrical; biarticulate palps present; antennae
hooks present. and tentacular cirri absent. Emergent notopodial spines
present.
Ancistrosyllis McIntosh 1879, A. groenlandica Mc-
Intosh 1879; 7 species. Pilargis Saint-Joseph 1899, P. verrucosa Saint-Joseph
Body flattened, with three antennae, biarticulate 1899; 7 species.
palps, two pairs of tentacular cirri. Peristomium dor- Body flattened; two antennae, biarticulate palps and
sally incised. Antennae shorter than palps. Emergent two pairs of tentacular cirri present. Emergent spines
notopodial hooks present. absent.
Cabira Webster 1879b, C. incerta Webster 1879b; Sigambra Muller 1858, S. grubii Muller 1858; 11
2 species. species.
Body cylindrical, with three antennae, biarticulate Body flattened, three antennae, biarticulate palps
palps and two pairs of tentacular cirri. Peristomium and two pa irs of tentacular cirri present. Antennae
dorsally incised. Dorsal and ventral cirri reduced, longer than palps. Emergent notopodial hooks present.
parapodia poorly developed. Emergent notopodial
Synelmis Chamberlin 1919c, S. simplex Chamberlin
hooks present.
1919c; 6 species.
Litocorsa Pearson, 1970, L. stremma Pearson 1970; Body cylindrical, with three antennae, biarticulate
only species. palps and two pairs of tentacular cirri present. Emergent
Body cylindrical without antennae and palps; two notopodial spines present.
pairs of tentacular cirri present. Emergent notopodial
spines present.
Loandalia Monro 1936, L. aberrans Monro 1936; Taxonomic Note
only species.
Talehsapia Fauvel 1932, with genotype T. annandalei
Body cylindrical with biarticulate palps; antennae
Fauvel 1932, has been considered a member of the
and tentacular cirri absent. Emergent spines absent.
family. As noted by Emerson and Fauchald (1971), it
Otopsis Ditlevsen 1917, O. longipes Ditlevsen 1917; cannot be considered a pilargiid, and has been con-
3 species. sidered an INCERTAE Sents. T. annandalei as reported by

Fauvel (1935) and Mesnil and Fauvel (1939) differ from represented in abyssal depths. In certain genera (Try-
the species as originally described; they are considered panosyllis, Autolytus and others), the structure of the
here as unidentifiable Parandalia spp. (Olga Hartman, trepan, the denticles along the cutting edge of the ever-
personal communication). sible pharynx, is of great importance. The pharynx is
only rarely eversed in preserved material. The exam-
ination of the trepan can be done through the body-wall,
Invalid Genera
if the specimen is very small and unpigmented. Larger
Glyphohesione Friedrich 1951, see Synelmis specimens, more than .5 mm across, or pigmented
Harpochaeta Korschelt 1893, see Ancistrosyllis specimens will have to be dissected. Examination of
Hermundura Muller 1858, indeterminable the eversible pharynx cannot be dispensed with in this
Kynephorus Ehlers 1920, see Synelmis family, even at the generic level, as is amply demon-
Phronia Webster 1879b, See Pilargis strated in the key below. Similarly, close examination
of the structure of the setae is also necessary, making
the identification of syllids a time-consuming occupa-
FAMILY SYLLIDAE GRUBE 1850
tion. Parapodia from anterior, median and posterior
Small to medium-sized nereidiform polychaetes region should be mounted on a slide for setal examina-
with usually, slender bodies (sometimes dorsoventrally tion under a compound microscope, and the whole
flattened). Three antennae and simple palps present, specimens should be scanned for modified setae of
the latter sometimes fused to each other. Two pairs any kind.
of tentacular cirri. Eversible pharynx armed with a Recent monographs include Imajima (1966a-d 1967)
single tooth or a circlet of smaller teeth or unarmed. who, in a series of papers revised the Japanese syllids.
Proventricle present in nearly all forms. Parapodia Gidholm has published a series of papers on the sub-
uniramous, dorsal cirri usually conspicuous, setae family Autolytinae (e.g. Gidholm 1962) and more
simple or composite. are expected. Hartmann-Schroder has also concentrated
The syllids are very common shallow-water forms, considerable attention on the syllids. Revision of the
and tend to be most numerous on hard substrates; California fauna is under way and may be expected
however, one sub-family, Exogoninae, also is well within a few years (Piltz, in preparation).
Key to Genera
Ia. Ventral cirri absent AUTOLYTINAE 2

lb. Ventral cirri present 9
2a (Ia). Dorsal cirri on first setiger only Procerastea
2b (Ia). Dorsal cirri on most setigers 3
3a (2b). All setae simple, occipital flap present Alluaudella
3b (2b). At least some setae composite, occipital flap absent 4
4a (3b). Dorsal cirri cylindrical 5
4b (3b). Dorsal cirri clavate or foliose 6
5a (4a). Bayonet-setae thick-shafted; segmental ciliary bands absent Proceraea
5b (4a). Bayonet-setae thin-shafted; ciliary bands present on every segment Autolytus
6a (4b). Eversible pharynx unarmed Phyllosyllis
6b (4b). Eversible pharynx with teeth 7
7a (6b). With projecting nuchal lobes, rather than flattened nuchal epaulettes Autosyllis
7b (6b). Nuchal organs flattened epaulettes 8
8a (7b). Antennae and dorsal cirri clavate Umbellisyllis
8b (7b). Antennae cylindrical, dorsal cirri flattened Myrianida
9a (Ib). Small forms (usually less than 8 mm)- palps fused for at least half their lengths ... EXOGONINAE
II
9b (1b). Larger forms, palps, if fused at all, only fused basally 10
l0a (9b). Palps fused at base; dorsal cirri smooth or irregularly wrinkled EUSYLLINAE 23
lob (9b). Palpi free to base or absent; dorsal cirri articulated SYLLINAE 43
1 Ia (9a). Antennae, tentacular and dorsal cirri absent Exogonella
I Ib (9a). At least tentacular cirri present, usually also antennae and dorsal cirri 12
12a (I lb). Two pairs of tentacular cirri present 13
12b (11b). One pair of tentacular cirri present 17


13a (12a). Antennae absent Exogonita
13b (12a). Three antennae present 14
14a (13b). Eversible pharynx non-muscular and sinuous Campesyllis
l 4b (13b). Eversible pharynx at least partially muscular, straight 15
15a (14b). Dorsal cirri long and filiform Brania
15b (14b). Dorsal cirri globular 16
16a (15b). Body with dorsal globular papillae in addition to the dorsal cirri Eurysyllis
I6b (15b). Body without dorsal globular papillae Plakosyllis
17a (12b). One antenna present; dorsal cirri rudimentary Spermosyllis
l7b (12b). Three antennae present; dorsal cirri well developed if sometimes short 18
1 8a (17b). Dorsal cirri long and slender 19
l8b (17b). Dorsal cirri short, ovoid or pyriform 20
19a (18a). Parapodia with erect, contractile dorsal lobes Anguillosyllis
l9b (18a). Parapodia without dorsal lobes Braniella
20a(I8b). Eversible pharynx unarmed 21
206 (18b). Eversible pharynx with a single anterior tooth 22
21 a (20a). Ventral cirri fused to lower edge of parapodia Exogonoides
21b (20a). Ventrial cirri free from parapodia Pseudexogone
22a (20b). Dorsal cirri pyriform (flask-shaped) Sphaerosyllis
22b (20b). Dorsal cirri papilliform or ovoid Exogone
23a (l0a). Eversible pharynx unarmed 25
23b (l0a). Eversible pharynx either with a single tooth or with a circlet of smaller teeth 24
24a (23b). Eversible pharynx with a single large tooth 30
24b (23b). Eversible pharynx with a series of teeth 41
25a (23a). Dorsum covered with small papillae Rhopalosyllis
25b (23a). Dorsum without papillae 26
26a (25b). Pharynx with an internal valve Pharyngeovalvata
26b (25b). Pharynx without an internal valve 27
27a (26b). Tentacular cirri absent; dorsal cirri rudimentary Fauvelia
27b (26b). Tentacular cirri present; dorsal cirri well developed 28
28a (27b). Antennae and tentacular cirri very short Atelesyllis
28b (27b). Antennae and tentacular cirri not noticeably short 29
29a (28b). Enlarged knobbed acicula present in anterior parapodia Streptosyllis
29b (28b). Enlarged knobbed acicula absent Syllides
30a (24a). Notacicula present Eudontosyllis
30b (24a). Notacicula absent 31
3Ia (30b). Antennae and dorsal cirri absent Nudisyllis
31b (30b). Antennae and dorsal cirri present 32
32a (31b). Three first segments fused with six pairs of tentacular cirri Irmula
32b (31b). Maximally two pairs of tentacular cirri on one segment 33
33a (32b). One pair of tentacular cirri present 34
33b (32b). Two pairs of tentacular cirri present 36
34a (33a). Dorsal cirri flattened Lamellisyllis
34b (33a). Dorsal cirri cylindrical 35
35a (34b). Composite setae spinigerous Parapionosyllis
35b (34b). Composite setae falcigerous Petitia
36a (33b). Tentacular and dorsal cirri very large, ovoid and inflated Clavisyllis
36b (33b). Tentacular and dorsal cirri more or less cylindrical 37
37a (36b). Middorsal tooth situated posterior in the eversible pharynx Opisthodonta
37b (36b). Middorsal tooth situated anterior in the eversible pharynx 38
38a (37b). Anterior margin of eversible pharynx denticulated 39
38b (37b). Anterior margin of eversible pharynx smooth 40
39a (38a). Median body-region with furcate, thick simple setae Synsyllis
39b (38a). Median body-region with composite falcigers Eusyllis
40a (38b). Parapodia long, palps twice as long as prostomium Dioplosyllis
40b (38b). Parapodia short and conical; palps maximally as long as prostomium Pionosyllis
41 a (24b). Eversible pharynx with a series of very small teeth Parautolytus

41 b (24b). Eversible pharynx with a limited number of large teeth 42

42a (41b). Body short, with few segments; large nuchal epaulettes present Amblyosyllis
42b (41b). Body longer, with numerous segments; nuchal epaulettes absent Odontosyllis
43a (lob). Palps absent; first segment with parapodia and setae Haplosyllides
43b (l0b). Palps present, first segment without parapodia and setae; usually with tentacular cirri 44
44a (43b). Eversible pharynx with a single tooth, a trepan or both 45
44b (43b). Eversible pharynx unarmed 46
45a (44a). Eversible pharynx with a trepan of several teeth; sometimes with an additional larger single tooth
as well 49
45b (44a). Eversible pharynx with a single tooth 50
46a (44b). Dorsum covered with small papillae Xenosyllis
46b (44b). Dorsum smooth 47
47a (46b). Parapodia with long, digitate distal lobes Branchiosyllis
47b (46b). Parapodia without distal lobes 48
48a (47b). Paired, posteriorly directed nuchal lappets present Parapterosyllis
48b (47b). Nuchal lappets absent Pseudosyllides
49a (45b). Palpi as long as prostomium; body cylindrical Geminosyllis
49b (45b). Palpi small and conical; body flattened Trypanosyllis
50a (45a). Middorsal tooth attached posteriorly Opisthosyllis
50b (45a). Middorsal tooth attached anteriorly 51
51a (50b). Setae simple, distally furcate or subdistally bossed Haplosyllis
51b (50b). At least some setae composite 52
52a (51 b). Dorsal cirri in posterior region alternately long and slender or large, bulbously fusiform
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Parasphaerosyllis
52b (51 b). Dorsal cirri may alternate between long and short, but all are of similar thickness 53
53a (52b). Tentacular and dorsal cirri with very few articles; dorsal cirri absent from second segment
. . . . . . .................................................................. Paratyposyllis
53b (52b). Tentacular and dorsal cirri with at least five articles; dorsal cirri present on second segment 54
54a (53b). Parapodia with pseudocomposite and simple setae in addition to the composite ones Syllis
54b (53b). Parapodia maximally with two simple setae in addition to the composite ones 55
55a (54b). Both composite spinigers and falcigers present Ehlersia
55b (54b). Only composite falcigers present Typosyllis
Generic Definitions EUSYLLINAE. Three antennae, palps separated

to the base. Antennae and tentacular cirri very short;
Alluaudella Gravier 1905c, A. madagascariensis Grav- large occipital flap present. Eversible pharynx unarmed.
ier 1905c; 2 species. Dorsal cirri emerge well above base of parapodia.
AUTOLYTINAE. Three short antennae and occipital
Autolytus Grube 1850, Nereis prolifera O.F. Muller
flap present. Palps completely fused; two pairs of
1788; 110 species.
tentacular cirri; setae all simple. Eversible pharynx
AUTOLYTINAE. Three antennae, two pairs of
unarmed.
tentacular cirri; first dorsal cirri longer than the rest.
Amblyosyllis Grube 1857, A. rhombeata Grube 1857; Nuchal epaulettes present. Eversible pharynx with
8 species. trepan with varying number of teeth. Each segment
EUSYLLINAE. Three long antennae, long palps with ciliated band. Bayonet-setae thin-shafted.
separated to the base. Dorsal cirri wrinkled. Paired
Autosyllis Imajima and Hartman 1964, A. japonica
long nuchal epaulettes present. Eversible pharynx
Imajima and Hartman 1964; only species.
with six or seven bi- tri- or pentacuspid teeth.
Anguillosyllis Day 1963, A. capensis Day 1963; only tentacular cirri; first dorsal cirri short. Dorsal cirri
species. clavate or foliose; nuchal projecting lobes present.
EXOGONINAE. Long slender forms with three
Branchiosyllis Ehlers 1887, B. oculata Ehlers 1887;
minute antennae; one pair of tentacular cirri; eversible
5 species.
pharynx unarmed. Parapodia with long contractile
SYLLINAE. Palps free to base; two pairs of ten-
dorsal lobes.
tacular cirri. Parapodia with long digitate distal lobes
Atelesyllis Pruvot 1930, A. rubrofasciata Pruvot 1930; attached pre- and postsetally. Eversible pharynx un-
only species. armed.

Brania Quatrefages 1866, Exogone pusilla Dujardin Eusyllis Malmgren 1867, E. blomstrandi Malmgren
EXOGONINAE. Two pairs of tentacular cirri; dorsal EUSYLLINAE. Three antennae and two pairs of
cirri long and filiform. Dorsal cirri longer than, ventral tentacular cirri. Eversible pharynx with middorsal tooth,
cirri as long as the setal lobes. Palpi as long as pros- margin denticulated. Occipital flap may be present.
tomium. Eversible pharynx with anterior dorsal tooth. Setae composite falcigers.
Braniella Hartman 1965, B. pupa Hartman 1965; 2 Exogone Orsted 1845, E. naidina Orsted 1845; 40
species. species.
EXOGONINAE. Three short, ovate antennae; one pair EXOGONINAE. Three antennae; one pair of ten-
of tentacular cirri; dorsal cirri long, slender and smooth, tacular cirri. Dorsal and ventral cirri shorter than setal
eversible pharynx unarmed. Composite spinigers. lobes; dorsal cirri ovoid or papilliform. Eversible
pharynx with a single tooth.
Campesyllis Chamberlin 1919a, C. minor Chamberlin
1919a; only species. Exogonella Hartman 1961, E. brunnea Hartman 1961;
EXOGONINAE. Three short antennae; two pairs of 2 species.
tentacular cirri. Eversible pharynx sinuous and non- EXOGONINAE. Antennae, tentacular and dorsal
muscular. cirri absent. Eversible pharynx with a single tooth.
Clavisyllis Knox 1957, C. anernata Knox 1957; only Exogonita Hartman and Fauchald 1971, E. oculata
species. Hartman and Fauchald 1971; only species.
EUSYLLINAE. Three antennae; two pairs of ten- EXOGONINAE. Antennae absent; two pairs of
tacular cirri; prominent nuchal epaulettes present. tentacular cirri present. Eversible pharynx with a single
Eversible pharynx smooth-rimmed with single dorsal tooth.
tooth. Tentacular and dorsal cirri large, ovoid and
inflated. Exogonoides Day 1963, E. antennata Day 1963; only
species.
Dioplosyllis Gidholm 1962, D. cirrosa Gidholm 1962; EXOGONINAE. Three ovoid antennae; one pair of
3 species. ovoid tentacular cirri. Dorsal cirri ovoid; ventral cirri
EUSYLLINAE. Three antennae, two pairs of ten- fused to parapodia. Eversible pharynx unarmed.
tacular cirri; nuchal ridges present or absent. Palps
very large and lingulate; parapodia long. Eversible Fauvelia Gravier 1900, F. martinensis Gravier 1900;
pharynx with middorsal tooth, smooth-rimmed or with only species.
a few teeth. EUSYLLINAE. Antennae absent; tentacular cirri
absent; dorsal cirri rudimentary. Eversible pharynx
Ehlersia Quatrefages 1865, Syllis sexoculata Ehlers unarmed.
1864; 15 species.
SYLLINAE. Three antennae and two pairs of ten- Geminosyllis Imajima 1966c, Trypanosyllis (Try-
tacular cirri; all anterior appendages articulated (except panedenta) ohma Imajima and Hartman 1964; only
palps). Eversible pharynx with middorsal tooth. Setae species.
include composite spinigers and falcigers and in poste- SYLLINAE. Body subcylindrical; three antennae,
rior setigers one or two simple setae per fascicle. two pairs of tentacular cirri. Eversible pharynx with
trepan of ten teeth and in addition a single large tooth.
Eudontosyllis Knox 1960, E. aciculata Knox 1960; All antennae and cirri slender and articulated.
only species.
EUSYLLINAE. Tentacular and dorsal cirri artic- Haplosyllides Augener 1922, H. floridana Augener
ulated, ventral cirri foliose. Eversible pharynx with 1922; only species.
middorsal tooth and denticulated margin; occipital SYLLINAE. Three antennae; palps absent. First
flap present. Notacicula present. segment with parapodia and setae. Dorsal cirri long,
ventral cirri short, all cirri smooth.
Eurysyllis Ehlers 1864, E. tuberculata Ehlers 1864;
3 species. Haplosyllis Langerhans 1879, Syllis spongicola Grube
EXOGONINAE. Body short and flattened. Three 1855; 10 species.
globular antennae; tentacular cirri and dorsal cirri also SYLLINAE. Three antennae, two pairs of tentacular
globular. Eversible pharynx with a trepan with 10 teeth cirri; all cirri articulated and slender. Eversible pharynx
and a middorsal tooth present. Dorsum covered with with single tooth. Setae simple, distally furcate or with
rows of globular papillae. a subdistal boss or knob.

Irmula Ehlers 1913, 1. spissipes Ehlers 1913; only Parasphaerosyllis Monro 1937b, P. indica Monro
species. 1937b; 4 species.
EUSYLLINAE. Three antennae; six pairs of ten- SYLLINAE. Three antennae; two pairs of tentacular
tacular cirri on three fused segments. All cirri smooth. cirri. Dorsal cirri anteriorly all slender and moniliform;
Eversible pharynx with anterior single tooth. posteriorly alternating between slender and large,
bulbously fusiform cirri. Eversible pharynx with mid-
Lamellisyllis Day 1960, L. comans Day 1960; only
dorsal tooth.
species.
EUSYLLINAE. Flattened, small form. Three foli- Pararyposyllis Hartmann-Schroder 1962b, P. paurocir-
aceous antennae; one pair of tentacular cirri. Eversible rata Hartmann-Schroder 1962; 2 species.
pharynx with single tooth. Dorsal cirri flattened; ventral SYLLINAE. Three antennae; two pairs of tentacular
cirri cylindrical. cirri; eversible pharynx with single tooth. All composite
setae falcigers; one or two simple setae present in each
Myrianida Milne Edwards 1845, Nereis pinnigera
of the posterior setigers. All antennae and cirri with
Montagu 1808; 8 species.
less than five articles; second segment without dorsal
AUTOLYTINAE. Three antennae; two pairs of ten-
cirri.
tacular cirri. All dorsal cirri flattened; antennae cylin-
drical. Eversible pharynx with trepan with varying Parautolytus Ehlers 1900, P. fasciatus Ehlers 1900;
numbers of teeth. 2 species.
EUSYLLINAE. Three antennae; two pairs of ten-
Nudisyllis Knox and Cameron 1970, N. tinihekia Knox
tacular cirri. Antennae and cirri smooth. Eversible
and Cameron 1970; only species.
pharynx finely denticulated, large tooth absent.
EUSYILLINAE. Antennae and dorsal cirri absent.
Eversible pharynx with a single tooth; margin smooth- Petitia Siewing 1955, P. amphophthalma Siewing
rimmed. 1955; only species.
EUSYLLINAE. Three antennae; one pair of ten-
Odontosyllis Claparede 1863, Syllis fuigurans Audouin
tacular cirri. Palpi biarticulate in adults. Eversible phar-
and Milne Edwards 1833a; 35 species.
ynx with a single tooth. Composite setae falcigerous.
tacular cirri. Occipital flap usually present. Eversible Pharyngeovalvata Day 1951, P. natalensis Day 1951;
pharynx with a series (less than 20) curved teeth. only species.
Opisthodonta Langerhans 1879, O. morena Langerhans
tacular cirri. Occipital flap present. Pharynx with
1879; 2 species.
valve; teeth absent.
tacular cirri; all antennae and cirri smooth. Eversible Phyllosyllis Ehlers 1897, P. albida Ehlers 1897, only
pharynx with a single large, posteriorly located tooth. species.
Some anterior parapodia with very thick acicula. AUTOLYTINAE. Three antennae; first segment
setose, with two large, foliose cirri. Eversible pharynx
Opisthosyllis Langerhans 1879, O. brunnea Langerhans
without teeth. Dorsal cirri foliose.
1879; 10 species.
SYLLINAE. Three antennae; two pairs of tentacular Pionosyllis Malmgren 1867, P. compacta Malmgren
cirri; occipital flap may be present. All antennae and 1867; 31 species.
cirri articulated. Eversible pharynx with posteriorly EUSYLLINAE. Three antennae; two pairs of ten-
attached mid-dorsal tooth; anterior margin smooth. tacular cirri. Tentacular and dorsal cirri smooth and
cylindrical. Eversible pharynx with single tooth; ante-
Parapionosyllis Fauvel 1923, Pionosyllis gestans
rior margin smooth.
Pierantoni 1903; 9 species.
EUSYLLINAE. Three antennae; one pair of ten- Plakosyllis Hartmann-Schroder 1956, P- brevipes
tacular cirri. Eversible pharynx with a single tooth. Hartmann-Schroder 1956; 2 species.
Composite setae spinigerous. EXOOONINAE. Body short, three globular antennae;
tentacular cirri and dorsal cirri also globular. Eversible
Parapterosyllis Hartmann-Schroder 1960a, P. sexocu-
pharynx with a trepan with ten teeth; a single tooth
lata Hartmann-Schroder 1960a; 2 species.
also present. Dorsal globular papillae absent.
SYLLINAE. Three antennae and two pairs of ten-
tacular cirri. Prostomium with paired posteriorly di- Proceraea Ehlers 1864, P. picta Ehlers 1864; 7 species.
rected nuchal lappets. All appendages articulated. AUTOLYTINAE. Three antennae; two pairs of ten-
Eversible pharynx unarmed. tacular cirri. Palps small and ventrally located. Eversible


pharynx with trepan with numerous teeth. Epaulettes Synsyllis Verrill 1900, S. longigularis Verrill 1900;
present. Bayonet-setae thick-shafted. Ciliary bands ab- 2 species.
sent from the setigers. EUSYLLINAE. Three antennae and two pairs of
Procerastea Langerhans 1884, P. nematodes Langer- tentacular cirri. Eversible pharynx with mid-dorsal
tooth; margin denticulated. Middle part of body with
hans 1884; 3 species.
l arge, distally furcate simple setae.
tentacular cirri. Dorsal cirri on first setiger only. Trypanosyllis Claparede 1864, Syllis zebra Grube
Eversible pharynx with trepan with numerous teeth. 1860; 26 species.
SYLLINAE. Body flattened with numerous short
Pseudexogone Augener 1922, P. backstromi Augener segments. Three antennae and two pairs of tentacular
1922; only species. cirri; all articulated. Trepan with several teeth; a single
EXOGONINAE. Three antennae;; one pair of ten- tooth may be present.
tacular cirri. Eversible pharynx unarmed. Dorsal cirri
present on second segment. Typosyllis Langerhans 1879, Syllis krohnii Ehlers 1864;
89 species.
Pseudosyllides Augener 1927b, P. curacaoensis Au- SYLLINAE. Three antennae and two pairs of ten-
gener 1927b; only species. tacular cirri; all articulated. Eversible pharynx with a
SYLLINAE. Three antennae; two pairs of tentacular single tooth. Setae include uni- or bidentate falcigers
cirri; antennae and cirri strongly articulated. Eversible in addition to one or two simple setae in each of the
pharynx unarmed with a smooth margin. posterior setigers.
Rhopalosyllis Augener 1913a, R. hamul(fera Augener Umbellisyllis Sars 1869, U. fasciata Sars 1869; 2
1913a; only species. species.
EUSYLLINAE. Three antennae and two pairs of AUTOLYTINAE. Three clavate antennae; two pairs
tentacular cirri. Antennae and cirri smooth. Dorsum of tentacular cirri. Nuchal organs foliaceous. Dorsal
covered with long papillae. Eversible pharynx unarmed. cirri clavate.
Spermosyllis Claparede 1864, S. torulosa Claparede Xenosyllis Marion and Bobretzky 1875, Syllis scabra
1864; 3 species. Ehlers 1864; 2 species.
EXOGONINAE. One antenna and one pair of ten- SYLLINAE. Three short, thick antennae; two pairs
tacular cirri. Eversible pharynx with a single tooth. of tentacular cirri. Tentacular cirri and dorsal cirri with
Dorsal cirri rudimentary, ventral cirri absent. few moniliform or collared articles. Dorsum covered
with small papillae. Eversible pharynx unarmed..
Sphaerosyllis Claparede 1863, S. hystrix Claparede
1863; 28 species. Taxonomic Notes
EXOGONINAE. Three antennae; one pair of ten-
tacular cirri. Dorsal cirri pyriform (flask-shaped), absent The subfamilies have been accepted strictly as de-
on second segment. Body with adhesive papillae. fined above; as a consequence several of the genera
have been moved from one subfamily to another. This
Streptosyllis Webster and Benedict 1884, S. arenae is not considered to be of any great importance: the
Webster and Benedict 1884; 7 species. differences between the subfamilies, especially between
EUSYLLINAE. Three antennae and two pairs of the EUSYLLINAE and SYLLINAE appear to be of
tentacular cirri. Eversible pharynx unarmed. Large more practical than scientific value.
knobbed acicula present in anterior setigers. The genus Irmula was originally described in the
Syllides Orsted 1845, S. longocirrata Orsted 1845; Hesionidae. It was moved to Syllidae by Day (1967).
13 species. It has a very isolated position in the family due to the
EUSYLLINAE. Three antennae and two pairs of presence of three modified anterior segments with six
tentacular cirri. Eversible pharynx unarmed; distal mar- pairs of tentacular cirri. However, the structure of
gin of pharynx smooth. No enlarged setae or acicula. pharynx, parapodia and setae is typically syllid, so
it appears best to retain it in the Syllidae.
Syllis Savigny 1818, S. monilaris Savigny 1818; 45 Hesiosyllis Wesenberg-Lund 1950, described as
species. intermediary between the syllids and the hesionids,
SYLLINAE. Three antennae; two pairs of tentacular is treated here among the latter, in that the structure
cirri, all articulated. Eversible pharynx with a single of the pharynx, setae, parapodia and tentacular cirri
tooth. Pseudocomposite and simple setae present in appear to resemble members of that family much more
addition to the composite setae in all parts of the body. than it resembles any member of the Syllidae.

Invalid Genera Protogrubea Czemiavsky 1881 a, see Brania

Pseudosyllides Czemiavsky 1882, see Amblyosyllis
Amytis Savigny 1818, see Proceraea Pterautolytus Ehlers 1907, see Autolytus
Anoplosyllis Claparede 1868, see Syllides Pterosyllis Claparede 1863, see Amblyosyllis
Aporosyllis Quatrefages 1865, see Syllis Sacconereis Muller 1853, see Autolytus
Autolytides Malaquin 1893, see Autolytus Salvatoria McIntosh 1885, see Brania
Brachysyllis Imajima and Hartman 1964, see Di-
Schmardia Quatrefages 1865, see Exogone
oplosyllis Stephanosyllis Claparede 1864, see Proceraea
Chaetosyllis Malmgren 1867, see Ehlersia Syllia Quatrefages 1865, see Exogone
Cirrosyllis Schmarda 1861, see Amblyosyllis and Sylline Grube 1860, see Autolytus
Autolytus
Sylline Claparede 1864, see Exogone
Claparedia Quatrefages 1865, see Eusyllis (?) Tetraglene Grube 1863, see Trypanosyllis
Crithida Gosse 1855, see Autolytus Thee Kinberg 1866b, see Typosyllis
Cystonereis K61liker in Koch 1846, see Exogone
Thylaciphorus Quatrefages 1865, see Amblyosyllis
Desmosyllis Verrill 1900, see Eusyllis
Trichosyllis Schmarda 1861, see Syllis
Diploceraea Grube 1850, see Autolytus
Virchowia Langerhans 1879, see Umbellisyllis
Doyeria Quatrefages in Milne Edwards 1848, NoMen
Xenosyllides Perejaslavzeva in Jakubova 1930, ques-
NUDUM tionably Umbellisyllis
Eucerastes Ehlers 1864, see Myrianida
Eurymedusa Kinberg 1866b, see Trypanosyllis
Exotokas Ehlers 1864, see Exogone FAMILY CALAMYZIDAE HARTMANN-SCHRODER 1971
Gattiola Johnston 1863, see Amblyosyllis
Gnathosyllis Schmarda 1861, see Syllis Body short, anteriorly and posteriorly rounded.
Gossia Quatrefages 1865, see Exogone (?) Prostomium small and without appendages with a suck-
Grubea Quatrefages 1865, see Brania ing mouth on the ventral side. All cirri (tentacular cirri,
Grubeosyllis Verrill 1900, see Brania dorsal and ventral cirri) short and digitate. Anal cirri
Hemisyllis Verrill 1900, see Haplosyllis absent. Setae composite. Eversible pharynx with stylet-
Hesperalia Chamberlin 1919a, ?Odontosyllis shaped sucking tube, otherwise without specializations.
Heterosyllis Claparede 1863, indeterminable The only known genus and species, Calamyzas
loda Johnston 1840, see Syllis amphictenicola Arwidsson 1932 is parasitic on the
Isosyllis Ehlers 1864, see Typosyllis ampharetid polychaete, Amphicteis gunneri (Sars 1835)
Lalage Miiller 1858, NoMEN NuDUM (see Syllis) from Sweden. A review can be found in Hartmann-
Lengerhansia Czemiavsky 1882, see Ehlersia Schroder 1971.
Laomedora Kinberg 1866b, indeterminable
Lapithas Kinberg 1866b, indeterminable
FAMILY NEREIDAE JOHNSTON 1845
Lophosyllis Sans 1867, indeterminable
Lycastis Savigny 1818, see Typosyllis Elongated, multi-segmented nereidiform polychaetes.
Microsyllis Claparede 1863, questionably Exogone Two, rarely one, antennae; palps biarticulated. Two
Monocerina Costa 1861a, indeterminable or four pairs of tentacular cirri. Eversible pharynx with
Nereisyllis Blainville 1828, see Syllis a pair of jaws and often accessory denticles or papillae.
Nicotia Costa 1864, see Amblyosyllis Parapodia nearly always biramous, usually with com-
Oophylax Ehlers 1864, see Exogone plex flattened lobes and cirri. Setae composite or sim-
Paedophylax Claparede 1868, see Exogone ple, spinigerous or falcigerous.
Pagenstecheria Quatrefages 1865, see Typosyllis The nereids are common forms in all depths, and
Parasitosyllis Potts 1 912, NoMEvNUDUM penetrate freshwater and to a very limited extent, even
Parexogone Mesnil and Caullery 1916, see Exogone terrestrial environments (Pflugfelder 1933). Some
Periboea Kinberg 1866b, indeterminable nereids are easy to maintain under laboratory conditions
Photocharis Ehrenberg 1835, indeterminable and have been used extensively for experimental re-
Platysyllis Grube 1878, indeterminable search. The most popular of the nereids thus used,
Podonereis Blainville 1818, see Autolytus is Hediste diversicolor, referred to in the experimental
Polybostrichus Orsted 1843a, see Proceraea literature incorrectly as Nereis diversicolor in most
Polymastus Claparede 1 864, see Eurysyllis cases; in other cases, more correctly as Neanthes di-
Polynice Savigny in Grube 1850, questionably Auto- versicolor.
lytus Some critical problems are associated with the recog-
Procorne Ehlers 1864, see Odontosyllis nition of species within the family, in that all species

cannot be recognized on strictly morphological charac- to be species-complexes, defined on non-morphological

ters (Smith 1958). It is suspected that in fact a series features. It is thus of importance that the provenance
of widespread species (Nereis pelagica, Neanthes of experimental organisms be stated clearly in all
virens, Platynereis dumerilii and others) may turn out publications.
Key to Genera
Ia. Peristomium forms a large ventral collar Cheilonereis
lb. Peristomium not ventrally enlarged 2
2a (lb). Some notopodia with pectinate branchiae 3
2b (I b). Branchiae absent 4
3a (2a). Branchiae arise from the dorsal cirrus; all setae composite spinigers Dendronereis
Frcuau 21. (A), Family NEREIDAE, Nereis vexillosa, Boiler Bay, Oregon, intertidal, dorsal view, 10x; (B), ventral view of the
above, 10x; (C) and (D), diagrams of the pharyngeal areas of nereids, in ventral and dorsal views; (E), median parapodium of
the above, 25x; (F), median parapodium of N. vexillosa from Dillon Beach, California, 50x.

3b (2a). Branchiae arise from the notopodial lobes; some composite falcigers present Dendronereides
4a (2b). Anterior ventrum with transverse fleshy ridges Australonereis
5
4b (2b). Anterior ventrum smooth
Sa (4b). Antennae absent 6
7
5b (4b). At least one antenna present
6a (5a). Two pairs of tentacular cirri present; anterior apodous segment absent Micronereis
6b (5a). Three pairs of tentacular cirri present; anterior apodous segment present Cryptonereis
7a (5b). A single median antenna present 8
7b (5b). Two antennae present 9
8a (7a). Paragnaths present on the maxillary ring; parapodia biramous (except the first two) Unanereis
8b (7a). Paragnaths absent; parapodia uniramous Dawbinia
9a (7b). Notocirri of parapodia 5-7 broadly elytraeform Kainonereis
9b (7b). Notocirri of parapodia 5-7 cylindrical and cirriform 10
10a (9b). Notopodia strongly reduced or absent 11
l0b (9b). Median and posterior notopodia well developed, with lobes and setae 14
I la (l0a). Notosetae present Namanereis
llb (l0a). Notosetae absent 12
12a(1lb). Tentacular cirri articulated Lycastoides
12b (lib). Tentacular cirri smooth 13
13a (12b). Notacicula present; antennae and cirri well developed Namalycastis
13b (12b). Notacicula absent; antennae and cirri reduced Lycastopsis
14a (l0b). Eversible pharynx with either papillae or paragnaths or both, in addition to the jaws 15
14b (l0b). Eversible pharynx with jaws, but otherwise smooth 16
15a (14a). Eversible pharynx with soft papillae only 20
15b (14a). Eversible pharynx with at least some paragnaths 25
16a (14b). Two pairs of tentacular cirri; apodous segment absent Micronereides
i 17
l6b (14b). Four pars of tentacular cirri; apodous segment present
17a (16b). Dorsal cirri attached basally on the notopodial superior lobes 18
17b (16b). Dorsal cirri attached distally on the notopodial superior lobes 19
18a (17a). Notopodial homogomph falcigers present in posterior setigers Rullierinereis
18b (17a). Notopodial homogomph falcigers absent Nicon
I9a (17b). Superior notopodial lobes long and straplike; inferior neuropodial lobe absent Steninonereis
19b (17b). Superior notopodial lobes large and foliose; inferior neuropodial lobes present Leptonereis
20a (15a). Pharyngeal papillae at least in part in tufts 21
20b (15a). Pharyngeal papillae solitary 22
21a (20a). All setae homogomph spinigers Tylonereis
21b (20a). Setae include also neuropodial homogomph falcigers in posterior setigers Laeonereis
22a (20b). Ventral cirri double at least in some setigers Ceratocephale
22b (20b). All ventral cirri simple 23
23a (22b). Accessory dorsal cirri on some anterior setigers; posterior dorsal cirri long and whiplike
........................................................................ Gymnonereis
23b (22b). Accessory dorsal cirri absent; posterior dorsal cirri not whiplike 24
24a (23b). Pharyngeal papillae on both rings; inferior neuropodial lobes absent; dorsal cirri distally attached ...
Tylorrhynchus
24b (23b). Pharyngeal papillae on oral ring only; inferior neuropodial lobes present; dorsal cirri basally at-
tached Kinberginereis
25a (15b). Eversible pharynx with both papillae and paragnaths Leonnates
25b (15b). Papillae absent, paragnaths present 26
26a (25b). Paragnaths present on one pharyngeal ring only 27
26b (25b). Paragnaths present on both pharyngeal rings 30
27a (26a). Paragnaths present on the maxillary ring only 28
27b (26a). Paragnaths present on the oral ring only 29
28a (27a). Paragnaths in eight groups, all rod-shaped Solomononereis
28b (27a). Paragnaths in patches and bands, all conical Ceratonereis
29a (27b). Notopodial homogomph falcigers present in posterior setigers Eunereis
29b (27b). Notopodial homogomph falcigers absent Websterinereis

30a (26b). All paragnaths conical 31

30b (26b). Cones and in addition either transverse or pectinate paragnaths or both present on the pharynx .. 35
31a (30a). All setae homogomph spinigers Nectoneanthes
31b (30a). At least some falcigers present 32
32a (31b). Middle and posterior neuropodia with single homogomph falcigers Hediste
32b (31 b). Homogomph falcigers, if present, in notopodial positions 33
33a (32b). With blunt simple falcigers in notopodia Cirronereis
33b (32b). Simple falcigers absent, composite falcigers present or absent 34
34a (33b). Notopodial homogomph falcigers present in posterior setigers Nereis
34b (33b). Notopodial homogomph falcigers absent Neanthes
35a (30b). Paragnaths include pectinate bars and usually small patches of cones; transverse smooth bars
absent Platynereis
35b (30b). Paragnaths include transverse smooth bars, patches of cones and sometimes pectinate bars 36
36a (35b). Superior notopodial lobes greatly expanded in posterior setigers; pectinate bars usually pres-
ent Pseudonereis
36b (35b). Superior notopodial lobes not expanded in any setigers; pectinate bars absent Perinereis
Generic Definitions Eversible pharynx with conical paragnaths on both

Australonereis Hartman 1954, Nereis (Leonnates) rings. Four pairs of tentacular cirri; biramous parapodia.
ehlersi Augener 1913a; only species. Notosetae include homogomph spinigers and blunt
simple falcigers; neurosetae heterogomph spinigers
Eversible pharynx with soft papillae on the maxillary
and falcigers.
ring, oral ring bare. Four pairs of tentacular cirri;
biramous parapodia. Notosetae homogomph spinigers; Cryptonereis Gibbs 1971, C. malaitae Gibbs 1971;
neurosetae homo- and heterogomph falcigers. With only species.
fleshy transverse ridges across anterior ventrum. Eversible pharynx without paragnaths or papillae.
Ceratocephale Malmgren 1867, C. loveni Malmgren Three pairs of tentacular cirri; uniramous parapodia.
1867; 11 species. Neurosetae heterogomph spinigers and falcigers. Fron-
Eversible pharynx with soft papillae on both rings. tal antennae absent; at maturity, parapodia biramous
with capillary setae.
Four pairs of tentacular cirri; biramous parapodia.
Setae include homogomph and heterogomph spinigers Dawbinia Benham 1950, D. aucklandica Benham
and heterogomph falcigers. Ventral cirri double on at 1950; only species.
lest some setigers, usually on most. Eversible pharynx without paragnaths or papillae.
Ceratonereis Kinberg 1866a, C. mirabilis Kinberg Two pairs of tentacular cirri; parapodia uniramous.
1866a; 53 species. Neurosetae homogomph spinigers and heterogomph
falcigers. A single median antenna present.
Eversible pharynx with paragnaths on the maxillary
ring only. Four pairs of tentacular cirri; biramous para- Dendronereides Southern 1921, D. heteropoda South-
podia. Notosetae include homogomph spinigers and ern 1921; 2 species.
falcigers; neurosetae homo- and heterogomph spinigers Eversible pharynx with soft papillae on both rings.
and heterogomph falcigers. Dorsal cirri attached basally Four pairs of tentacular cirri; parapodia biramous.
to the superior notopodial lobe; inferior neuropodial Branchiae present as a subdivision of the notopodial
lobe may be present. superior lobes. Neuropodial inferior lobes absent.
Cheilonereis Benham 1916, Nereis cyclurus Harrington Notosetae homogomph spinigers; neurosetae home-
1897; 2 species. and heterogomph spinigers and heterogomph falcigers.
Eversible pharynx with conical paragnaths on both Dendronereis Peters 1854, D. arborifera Peters 1854;
i
rings. Four pairs of tentacular cirri; b ramous parapodia. 3 species.
Notosetae homogomph spinigers; neurosetae homo- Eversible pharynx smooth or with soft papillae. Four
gomph spinigers and heterogomph falcigers. Superior pairs of tentacular cirri; parapodia biramous. Branchiae
notopodial lobes large and foliose with dorsal cirrus present as subdivisions of the dorsal cirri. Neuropodia
attached medially. Peristomium greatly expanded in anterior setigers multifid. All setae homogomph
ventrally. spinigers.
Cirronereis Kinberg 1866a, C. gracilis Kinberg 1866a; Eunereis Mahngren 1867, Nereis longissima Johnston
only species. 1840; 7 species.

Eversible pharynx with paragnaths on oral ring only. Superior notopodial lobes large and foliose in posterior
Four pairs of tentacular cirri; parapodia biramous. setigers. Notosetae homogomph spinigers; neurosetae
Notosetae homogomph spinigers and falcigers; neuro- heterogomph spinigers and falcigers, the latter with
setae homo- and heterogomph spinigers and hetero- long appendages. Inferior neuropodial lobe present.
gomph falcigers.
Lycastoides Johnson 1903, L. alticola Johnson 1903;
Gymnonereis Horst 1919a, Gymnorhynchus sibogae only species.
Horst 1918; 2 species. Eversible pharynx without papillae or paragnaths.
Eversible pharynx with papillae on the oral ring only. Four pairs of tentacular cirri present; parapodia uni-
Four pairs of tentacular cirri; parapodia biramous. Ac- ramous. Neurosetae heterogomph falcigers and spin-
cessory dorsal cirri present on some anterior segments; igers. Tentacular cirri jointed; eyes absent.
posterior dorsal cirri long and whiplike. All setae Lycastopsis Augener 1922, L. beameri Augener 1922;
homogomph or slightly hemigomph spinigers. 6 species.
Hediste Malmgren 1867, Nereis diversicolor O.F. Eversible pharynx without papillae or paragnaths.
Maller 1776; only species. Three pairs of tentacular cirri present; parapodia uni-
Eversible pharynx with conical paragnaths on both ramous. Neurosetae heterogomph spinigers and falcigers.
rings. Four pairs of tentacular cirri; parapodia biramous. Antennae and cirri reduced.
Notosetae homogomph spinigers. Neurosetae homo- Micronereides Day 1963, M. capensis Day 1963;
and heterogomph spinigers; heterogomph falcigers. A only species.
single homogomph falciger present in median and Eversible pharynx without papillae or paragnaths.
posterior neuropodia. Two pairs of tentacular cirri present; parapodia bi-
Kainonereis Chamberlin 1919c, K. alata Chamberlin ramous. Apodous segment absent. All setae homo-
1919c; only species. gomph spinigers.
Eversible pharynx without paragnaths or papillae. Micronereis Claparede 1863, M. variegata Claparede
Four pairs of tentacular cirri present; parapodia bi- 1863; 5 species.
ramous. Antennae bifid at the tips; broad elytraeform Eversible pharynx without papillae or paragnaths.
dorsal cirri on parapodia 5-7. Two pairs of tentacular cirri present; parapodia bi-
Kinberginereis Pettibone 1971a, Nereis (Leptonereis) ramous. Apodous segment absent. Antennae absent.
inermis Hoagland 1920; only species. All setae homogomph spinigers.
Eversible pharynx with soft papillae on the oral ring Namalycastis Hartman 1959, Paranereis abiuma Maller
only. Four pairs of tentacular cirri; parapodia biramous. in Gmbe 1871; 18 species.
Notosetae homogomph spinigers; neurosetae homo- Eversible pharynx without papillae or paragnaths.
gomph and heterogomph spinigers. Inferior neuro- Four pairs of tentacular cirri; parapodia sub-biramous
podial lobe present; dorsal cirri basal. or uniramous. Notosetae usually absent; neurosetae
Laeonereis Hartman 1945, Nereis culveri Webster heterogomph spinigers and falcigers. Neuropodia with
1879a; 6 species. a single setal lobe only. Notopodial superior lobes
Eversible pharynx with tufts of papillae on both prolonged in posterior setigers.
rings and large solitary papillae on area VI. Four pairs Namanereis Chamberlin 1919c, Lycastis quadraticeps
of tentacular cirri; parapodia biramous. Notosetae Blanchard 1849; 2 species.
homogomph spinigers; neurosetae homogomph spinigers Eversible pharynx smooth or with soft papillae.
and falcigers, the latter in posterior setigers. Inferior Three or four pairs of tentacular cirri present; parapodia
neuropodial lobes present. biramous with notopodia strongly reduced. Neurosetae
include heterogomph spinigers and falcigers,
Leonnates Kinberg 1866a, L. indicus Kinberg 1866a;
10 species. Neanthes Kinberg 1866a, N. vaalii Kinberg 1866a;
Eversible pharynx with papillae on the oral ring 50 species.
and paragnaths on the maxillary ring. Four pairs of Eversible pharynx with conical paragnaths on both
tentacular cirri; parapodia biramous. Notosetae homo- rings. Four pairs of tentacular cirri, parapodia biramous.
gomph spinigers; neurosetae heterogomph falcigers Notosetae homogomph spinigers; neurosetae homo- and
with coarsely serrated blades. heterogomph spinigers and heterogomph falcigers.
Leptonereis Kinberg 1866a, L. laevis Kinberg 1866a; Nectoneanthes Imajima 1972, Nereis (Alitta) oxypoda
2 species. Marenzeller 1879; 2 species.
Eversible pharynx without papillae and paragnaths. Eversible pharynx with conical paragnaths on both
Four pairs of tentacular cirri, parapodia biramous. rings. Four pairs of tentacular cirri present; parapodia


biramous. All setae homogomph spinigers. Expanded Eversible pharynx with eight groups of rod-shaped
superior notopodial lobes in median and posterior paragnaths on the maxillary ring; oral ring bare. Four
setiger, with dorsal cirrus inserted between the superior pairs of tentacular cirri; parapodia biramous. Notosetae
and the median lobes. include homogomph spinigers and falcigers; neuro-
Nereis Linnaeus 1758, N. pelagica Linnaeus 1758; setae hemi-, heterogomph spinigers and heterogomph
134 species. falcigers.
Eversible pharynx with conical paragnaths on both Steninonereis Wesenberg-Lund 1958, S. martini
rings. Four pairs of tentacular cirri present; parapodia Wesenberg-Lund 1958; only species.
biramous. Notosetae include homogomph spinigers Eversible pharynx without papillae and paragnaths.
and falcigers, the latter in median and posterior setigers; Four pairs of tentacular cirri; parapodia biramous.
neurosetae include homo- and heterogomph spinigers Notosetae homogomph spinigers; neurosetae home-
and heterogomph falcigers. and heterogomph spinigers, and heterogomph falcigers.
Nicon Kinberg 1866a, N. pictus Kinberg 1866a; 15 Superior notopodial lobes, long and straplike in poste-
rior setigers.
species.
Eversible pharynx without papillae or paragnaths. Tylonereis Fauvel 1911, T. bogoyawlenskyi Fauvel
Four pairs of tentacular cirri; parapodia biramous. Noto- 1911; 2 species.
setae homogomph spinigers; neurosetae homogomph Eversible pharynx with soft papillae on the maxillary
spinigers and falcigers. Inferior neuropodial lobes ring only. Four pairs of tentacular cirri present; para-
present. podia biramous. All setae homogomph spinigers.
Perinereis Kinberg 1866a, P. novaehollandiae Kinberg Tylorrhynchus Grube 1869a, Nereis heterochaeta Qua-
1866a; 60 species. trefages 1865; 2 species.
Eversible pharynx with conical and transverse para- Eversible pharynx with soft papillae on both rings.
gnaths on both rings; four pairs of tentacular cirri; Four pairs of tentacular cirri; parapodia biramous.
parapodia biramous. Notosetae homogomph spinigers; Notosetae hemigomph spinigers; neurosetae hemi- and
neurosetae homo- and heterogomph spinigers and heterogomph spinigers and heterogomph falcigers.
heterogomph falcigers. Inferior neuropodial lobes absent.
Plasynereis Kinberg 1866a, P. magalhaensis Kinberg Unanereis Day 1962, U. macgregori Day 1962; only
1866a; 20 species. species.
Eversible pharynx with paragnaths on both rings, Eversible pharynx with conical paragnaths on the
including cones, and pectinate bars. Four pairs of maxillary ring. Four pairs of tentacular cirri; parapodia
tentacular cirri; parapodia biramous. Notosetae homo- biramous. Notosetae homogomph spinigers and falcigers;
gomph spinigers and falcigers, the latter sometimes neurosetae homo- and heterogomph spinigers and
fused to form simple falcigers; neurosetae include heterogomph falcigers. A single median antenna
homo- and heterogomph spinigers and heterogomph present.
falcigers.
Websterinereis Pettibone 1971a, Nereis tridentata
Pseudonereis Kinberg 1866a, P. gallapagensis Kinberg Webster 1880; only species.
1866a; 7 species. Eversible pharynx with paragnaths on the oral ring
Eversible pharynx with paragnaths on both rings, only. Four pairs of tentacular cirri; parapodia biramous.
including cones, transverse smooth bars and pectinate Notosetae homogomph spinigers; neurosetae homo-
bars. Four pairs of tentacular cirri; parapodia biramous. and heterogomph spinigers and heterogomph falcigers.
Notosetae include homogomph spinigers and falcigers;
neurosetae homo- and heterogomph spinigers and
heterogomph falcigers. Taxonomic Notes
The generic subdivision of the nereids has been
Rullierinereis Pettibone 1971a, Leptonereis zebra
based mainly on the pharyngeal structures and the
Rullier 1963; 5 species.
presence of specific kinds of setae in the parapodial
Eversible pharynx without papillae or paragnaths.
rami. Both characters are subject to some variation
Four pairs of tentacular cirri; parapodia biramous.
Notosetae include homogomph spinigers and falcigers, within each genus, and especially the pharyngeal struc-
tures require accurate dissection of the anterior end.
the latter in posterior setigers; neurosetae homo- and
Pettibone (1971a) introduced characters of the para-
heterogomph spinigers and heterogomph falcigers.
podial lobes (called ligules by Pettibone) as major fea-
Inferior neuropodial lobes present.
tures in the generic identification; this may be valid,
Solomononereis Gibbs 1971, S. maranensis Gibbs but the character has the distinct drawback that it is
1971; only species. dependent on interpretation of shapes, which is noto-

riously dependent on the experience of the observer, Pisenoe Kinberg 1866a, see Platynereis
and very difficult to quantify. Pettibone in the same Podonereis Blainville 1826, indeterminable
paper lumped series of species based on overlapping Praxithea Malmgren 1867, see Nereis
ranges in different characters; this is unfortunate, since Protolycoris Hatschek 1893, NomEANuouM
the variability within any single population of these Stratonice Malmgren 1867, see Perinereis
animals has never been examined and quantified in de- Tetratrocha Sveshnikov 1959, larvae, no species named
tail. The generic key given above, reflects the added Thoosa Kinberg 1866a, see Nereis
insights of Pettibone at the generic level, but the num- Typhlonereis Hansen 1878, indeterminable
bers of species indicated for each genus is higher than as Uncinereis Chamberlin 1919c, see Platynereis
given by Pettibone, reflecting the more conservative
approach taken to lumping at the specific level in this
study. FAMILY Antonbruunidae NEW NAME
Nereidiform polychaetes with cylindrical bodies,

Invalid Genera three antennae and simple palps. Eversible pharynx
unarmed. Two pairs of tentacular cirri present. Para-
Aceronereis Blainville 1818, NOMEN NUDUM podia biramous, but notopodia reduced to dorsal cirri
Alitta Kinberg in Malmgren 1865, see Neanther supported by internal acicula. All setae simple. Sexual
Arete Kinberg 1866a, see Perinereis dimorphism present, with small males; inquilines in
Branchionereis Blainville 1818, NOMENNDDUM bivalves.
Chaunorhynchus Chamberlin 1919c, we Ceratocephale This family is known for a single genus, Antonbruunia
Chinonereis Chamberlin 1924, see Tylorrhynchus Hartman and Boss 1965 with genotype A. viridis living
Cirroceros Claparede 1863, indeterminable in the mantle cavity of the bivalve Lucina fosteri Hart-
Cirronereis Blainville 1818, indeterminable man and Boss.
Gnatholycastis Ehlers 1920, see Perinereis
Gymnorhynchus Horst 1918, see Gymnonereis
Hedyle Malmgren 1867, see Perinereis Suborder Glyceriformia
Heminereis Quatrefages 1865, indeterminable
Heteronereis Orsted 1843a, see Nereis Phyllodocida with two pairs of antennae; palps and
Iphinereis Malmgren 1865, see Platynereis tentacular cirri absent. Proboscis either unarmed, with
Leontis Malmgren 1867, see Platynereis four jaws or with a circlet of jaws. First parapodia
Lepidonereis Blainville 1818, indeterminable lateral.
Lipephila Malmgren 1867, see Perinereis
Lycastella Feuerbom 1932, see Namanereis
Lycastis Audouin and Milne Edwards 1833b, see FAMILY GLYCERIDAE GRUBE 1850
Namalycastis
Lycastoides Jakubova 1930, see Namanereis Glyceriform polychaetes with long, slender bodies
Lycoris Savigny 1818, see Nereis and conical prostomia. Eversible pharynx with four
Mastigonereis Schmarda 1861, see Pseudonereis jaws in a cross. Parapodia either all biramous or all
Meganereis Blainville 1818, indeterminable uniramous. Neurosetae composite; notosetae, when
Naumachius Kinberg 1866a, see Pseudonereis present, simple.
Nectonereis Verrill 1873b, see Platynereis The glycerids are long, slender polychaetes with
Nereilepas Blainville 1828, see Neanthes numerous segments. Perhaps their most noticeable
Nicomedes Kinberg 1866a, indeterminable feature is the very long, cylindrical eversible pharynx,
Nossis Kinberg 1866a, indeterminable which they will evert when disturbed. At the tip of
Paralycastis Ehlers 1920, see Perinereis the pharynx are four short, usually black jaws. Glyc-
Paranereis Kinberg 1866a, see Pseudonereis erids are present, mainly in soft sandy or muddy sub-
Phaetusa Castelnau 1842, see Hediste strates, in all depths; they are rarely present in large
Phyllonereis Hansen 1882, see Pseudonereis numbers. They appear to be mainly carnivores.
Key to Genera
Ia. Parapodia uniramous throughout Hemipodus

lb. Parapodia biramous throughout 2
2a (lb). Prostomium long, with more than three rings; aileron of jaws with lateral wing Glycera
2b (lb. Prostomium short, with about three rings; aileron of jaws a simple rod Glycerella

Prostomium long, with seven to ten vague annuli;

aileron of jaws rod-shaped. Pharyngeal organs elongate
oval or filamentous. Parapodia uniramous; all setae
composite spinigers.
Taxonomic Notes
The generic subdivision, essentially in two major
genera, has been stable for the last 50 years and not
much change is anticipated. Identification of species,
especially in certain groups of Glycera is quite dif-
ficult, and the number of species may be subject to
considerable adjustment. Specific identification de-
pends on study of the pharyngeal organs, in
addition to parapodial lobes and branchiae. Pharyngeal
organs can be characterized only by very close work
under compound microscopes. It is of great importance
that the light be very accurately adjusted, since the
characteristic ridges may be very difficult to see and
depend on the full resolution of the microscope, not
because of their small sizes, but because of their
structure. Branchiae may be retractable, and frequently
are retracted in parts of the body; the whole body
must be scanned for the presence or absence of these
structures.
B Invalid Genera
FICOan 22. (A) Family GLYCERIDAE, Glycera americana,
off Santa Barbara, intertidal, 10x; (B), Glycera capitata, Euglycera Verrill 1881, see Glycera
median parapodium, posterior view, after Hartman, 1950, 64x. Hamiglycera Ehlers 1908, see Glycera
Hemipodua Kinberg 1866b, see Hemipodus
Proboscidea Blainville 1825, see Glycera
Rhynchobolus Claparede 1868, see Glycera
Telake Chamberlin 1919c, see Glycera
Generic Definitions
Glycera Savigny 1818, G. unicornis Savigny 1818; FAMILY GONIADIDAE KINBERG 1866b
55 species. Glyceriforms with long and slender bodies. Pros-
Prostomium long, with at least three, usually five to tomium is conical; eversible pharynx with a circlet of
seven annuli. Aileron of jaw with lateral wing. Pharyn- smaller and larger jaw-pieces. Parapodia anteriorly
geal organs of many kinds. Notosetae simple, capillary uniramous, posteriorly biramous, rarely all uniramous.
or acicular; neurosetae composite spinigers. Neurosetae composite, notosetae simple.
Glycerella Arwidsson 1899, Hemipodus magellanicus The goniadids often are considered as part of the
McIntosh 1885; 2 species. glycerids, but the two groups differ sufficiently that
Prostomium short, with maximally four annuli. a recognition at the family level is warranted. They
Aileron of jaws rodlike. Pharyngeal organs long and resemble the glycerids in that their most remarkable
slender. Notosetae capillaries; neurosetae composite structure is the eversible pharynx, covered with pharyn-
spinigers. geal organs, very long, usually slender and crowned
with a series of teeth. The pharyngeal organs are con-
Hemipodus Quatrefages 1865, Glycera rosea Blain- siderably larger than in the glycerids, and are partly
ville in Quatrefages 1865, 15 species. sclerotinized in some genera (Glycinde, Bathyglycinde).
Key to Genera
Ia. All parapodia uniramous Progoniada
l b. Anterior parapodia uniramous, posterior ones biramous 2
2a (lb). Eversible pharynx with organs of many kinds 3

2b (lb). Eversible pharynx with organs of one or a few kinds 4

3a (2a). Notosetae slender capillaries Bathyglycinde
3b (2a). Notosetae knobbed or falcate hooded hooks Glycinde
4a (2b). Eversible pharynx with chevrons 5
4b (2b). Eversible pharynx without chevrons 6
5a (4a). Neuropodia with spinigers only Goniada
5b (4a). Neuropodia with both spinigers and falcigers Goniadella
6a (4b). Neuropodia with spinigers only Ophioglycera
6b (4b). At least some falcigers present 7
7a (6b). Notopodia reduced to a rounded knob with a few thick acicular setae and a citrus Goniadides
7b (6b). Notopodia well developed 8
8a (7b). Anterior setae all falcigers; posterior ones both spinigers and falcigers Bookhoutia
8b (7b). Anterior setae all falcigers, posterior ones all spinigers Goniadopsis
Generic Definitions Posterior segments biramous. Chevrons absent.

Neuropodia in unimmous part of body with composite
Bathyglycinde Fauchald 1972, B. mexicana Fauchald falcigers; those in biramous part, spinigers. Notosetae
1972; 3 species. acicular spines.
Posterior segments biramous. Chevrons absent,
pharyngeal organs large and of several kinds. Notosetae
capillaries; neurosetae composite spinigers.
Bookhoutia Mohammad 1973, B. oligognatha Moham-
mad 1973; only species.
Posterior segments biramous. Chevrons absent;
e
pharyngeal organs small and of one kind. Notosetae
acicular; anterior neurosetae all falcigers, posterior -VoMr-
neurosetae both falcigers and spinigers.
Glycinde Mu ller 1858, G. multidens Muller 1858; 20
,I
species.
Posterior segments biramous. Chevrons absent;
pharyngeal organs large and of several kinds. Noto-
setae knobbed or falcate hooded hooks; neurosetae
composite spinigers.
Goniada Audouin and Milne Edwards 1833b, G.

emerita Audouin and Milne Edwards 1833b; 34
species.
Posterior segments biramous. Chevrons present;
pharyngeal organs small, mainly of one kind. Notosetae
acicular or capillary; neurosetae composite spiningers.
Goniadella Hartman 1950, Eone gracilis Verrill 1873b;
2 species.
Posterior segments biramous. Chevrons present;
pharyngeal organs of one kind. Composite spinigers
and falcigers present on all neuropodia. s
r
Goniadides Hartmann-Schroder 1960a, G. aciculata

Hartmann-Schroder 1960a; 2 species.
Posterior segments biramous. Chevrons absent. Noto-
podia reduced to short rounded lobes with a cirrus and A
one or two coarse acicular setae. All neurosetae com- FIGURE 23. (A), Family GONIADIDAE, Glycinde armigera,
posite falcigers. anterior end with pharynx half everted, after Hartman, 1950,
12.5x; (B), Goniada brunnea, median parapodiurn, posterior
Goniadopsis Fauvel 1928a, G. agnesiae Fauvel 1928a; view, after Hartman, 1950, 31x; (C), anterior end of the above,
3 species. 25x.

Ophioglycera Verrill 1885, O. gigantea Verrill 1885; The lacydoniids somewhat resembles the nephtyids,
6 species. in that both noto- and neuropodia are developed equally,
Posterior segments biramous. Chevrons absent, but the parapodia and setal structures, as well as the
pharyngeal organs of one kind and short. Notosetae structure of the prostomium, ally them more closely
slender and acicular; all neurosetae composite spinigers. with the glycerids than with any other group of poly-
Progoniada Hartman 1965, P. regularis Hartman 1965; chaetes. They traditionally have been considered as
an appendix to the phyllodociform polychaetes, mainly
2 species.
All segments uniramous. Chevrons present. Neuro- because of the lack of proboscideal armament. The
setae include composite falcigers and spinigers in all lacydoniids recently were reviewed by Uschakov (1972);
parapodia. the present treatment follows his closely.
Taxonomic Notes
The genera appear well established in the Goniadidae.

The only difficult point in the key above appears in
dichotomy 2 on the question of a few or many kinds
of pharyngeal organs. In Glycinde and Bathyglycinde
at least three distinct kinds of organs are present; in
most species at least two kinds are distinctly chitinized
projections. In the other genera, most species have
only one kind of pharyngeal organ, but in some cases
two kinds may be identifiable.
Invalid Genera
Eone Malmgren 1866, see Glycinde
Epicaste Kinberg 1866b, see Glycinde
Lacharis Kinberg 1866b, indeterminable
Leonnatus Kinberg 1866b, see Goniada
FAMILY LACYDONIIDAE BERGSTROM 1914

Glyceriform polychaetes with short and slender
bodies; prostomium trapezoidal in outline; eversible
pharynx unarmed. First parapodia uni- or biramous, FIGuRe 24. (A), Family LACYDONIIDAE, Paralacydonia
all others biramous. Neurosetae composite, notosetae sp., San Clemente Basin, California, 1500 m, 25x; (B), median
simple. parapodium of the above, 50x.
Key to Genera
la. Tentacular cirri present Lacydonia
lb. Tentacular cirri absent 2
2a (lb). Antennae short and biarticulated; first setiger uniramous Paralacydonia
2b (I b). Antennae long and smooth; first setiger biramous Pseudolacydonia
Generic Definitions Pseudolacydonia Rullier 1965b, P. caeca Rullier

Lacydonia Marion and Bobretzky 1875, L. miranda 1965b; only species.
Marion and Bobretzky 1875; 4 species. Antennae long and slender, not articulated; tentacular
Antennae short and smooth; tentacular cirri present; cirri absent; all parapodia biramous.
first two-three parapodia uniramous.
Paralacydonia Fauvel 1913, P. paradoxa Fauvel 1913; The following five families do not appear to be
3 species. closely related to any other Phyllodocida nor are they
Antennae short and biarticulated; tentacular cirri obviously related to each other. They are listed in
absent; only first parapodia uniramous. alphabetical order.

The three families with pelagic members, Iospilidae, The iospilids often are considered allied with the
Tomopteridae and Typhloscolecidae, usually are con- phyllodocids, but appear to differ rather sharply from
sidered with the bulk of the other pelagic polychaetes, members of this family in most of the characters usually
most of which are related to the Phyllodocidae. This considered at the familial level. It is here considered
connection seems based on adaptive convergencies a member of the order Phyllodocida, but has not been
to the pelagic environment, such as a frequently foliose assigned to any suborder or super-familial group. The
condition of the parapodial lobes; a reduction in the family was revised by Dales and Peter (1972) and
number and importance of the setae and the lightly Uschakov (1972).
built, often translucent bodies in these forms.
The nephtyids and sphaerodorids either are con-
sidered related to the glyceriforms or placed in the
vicinity of the nereids. The nephtyids are extremely
poorly cephalized compared to most other polychaetes,
in that even the first setiger carries small, but recog-
nizable parapodia and setae. The lack of cephalization
places the nephtyids close to the phyllodocids, from
which they differ sharply in the development of the
parapodia, in that they have one pair of antennae and
one pair of palps, according to the innervation pattern
whereas the phyllodocids have at least two pairs of
antennae and true palps are absent. For these reasons,
it appears for the time being best to leave the nephtyids
as a free-standing family within the order Phyllodocida.
The sphaerodorids, however, appear to have an exten-
sive and varied degree of cephalization (Fauchald
1974b) and appear isolated in the Phyllodocida. They
are not related to the nephtyids. The iospilids differ
from the phyllodociform families in the presence of
palps and the lack of antennae. The same can be said
for the tomopterids and the typhloscolecids (Uschakov
1972).
In all five cases, the presence and structure of the
anterior appendages (sometimes the lack of such ap-
pendages), makes it difficult to ally these forms with
any other polychaetes. Rather than forcing the issue,
and insist that all families must be allied with a sub-
order, it here is considered best to leave all five as
separate entities, without any commitment as to further
affiliation.
FAMILY IOSPILIDAE BERGSTROM 1914
Pelagic, small, slender Phyllodocida. Prostomium

without antennae; two short palps present. Parapodia /
uniramous, all setae composite. Eversible pharynx FIGURE 25. Family ISOPILIDAE, Phalacrophorus pictus
present, armed in some cases. borealis, modified after Uschakov, 1972, 50x.
Key to Genera
Ia. Eversible pharynx with a pair of curved jaws Phalacrophorus

lb. Eversible pharynx unarmed 2
2a (lb). Up to 11 anterior segments with reduced parapodia lospilopsis
2b (lb). Parapodia fully developed from segment 5 3
3a (2b). Dorsal and ventral cirri present, but small on segments 2-3 Paraiospilus
3b (2b). Dorsal and ventral cirri absent on segments 2-3 lospilus

Generic Definitions Maximally ten anterior segments with reduced para-

podia. Eversible pharynx with paired jaws.
lospilopsis Augener 1922, 1. antillensis Augener 1922;
only species. FAMILY NEPHTYIDAE GRUBE 1850
Upp to 11 anterior segments with rudimentary para-
podia; jaws absent. Phyllodocida with long, slender bodies with
quadrangular cross-sections. One pair of antennae and
lospilus Viguier 1886, 1. phalacroides Viguier 1886; one pair of very short palps present. A pair of lateral
2 species. jaws present. Parapodia biramous with both rami well
Maximally four anterior segments with reduced developed with complex pre- and postsetal lobes. All
parapodia; dorsal and ventral cirri absent on segments setae simple. Interramal, respiratory cirri present in
2 and 3; jaws absent. nearly all forms.
Paraiospilus Viguier 1911, P. affinis Viguier 1911;
The nepthyids may superficially be confused with
only species. the sigalionids in that both groups have long, straight-
Maximally four anterior segments with reduced sided bodies abruptly tapering anteriorly, and rather
parapodia; dorsal and ventral cirri present, but rudi- more gently posteriorly. However, the nephtyids lack
mentary on segments 2 and 3; jaws absent. obvious, long appendages anteriorly, and of course
the scales present in the sigalionids. Nephtyids are
Phalacrophorus Greeff 1879, P. pictus Greeff 1879; present at all depths, and are most common in sandy
3 species. and muddy substrates.
FIGURE 26. (A), Family NEPHTYIDAE, Nephtys californiensis, off Santa Catalina Island, California, 50 m, 15x; (B), median
parapodium of the above, 15x; (C), Family SPHAERODORIDAE, Sphaerodoropsis sphaerulifer, off Santa Catalina Island,
70 m, 75x; (D), Family TOMOPTERIDAE,Tonwpteris sp., off central California, pelagic, 5x; (E), parapodium of the above, 10x.

Key to Genera
la. Interramal cirri rudimentary or absent Micronephthys

l b. Interramal cirri well developed, recurved or involute 2
2a (lb). Interramal cirri recurved Nephtys
2b (Ib). Interramal cirri involute 3
3a (2b). Eversible pharynx with subterminal and terminal papillae Aglaophamus
3b (2b). Eversible pharynx without subterminal and terminal papillae Inermonephtys
Generic Definitions The first start of the interrama cirri is seen best from
the lateral side, and a probe must be used to lift the
Aglaophamus Kinberg 1866b, A. lyratus Kinberg
notopodial cirrus so that the small, barely emerging
I866b; 45 species.
interramal cirrus can be seen on its ventral side. Recent
Eversible pharynx with 14 (rarely 16) rows of pa-
revisions include Hartman (1950) and Fauchald
pillae. Interramal cirri involute; acicula distally hooked
(1968a).
in most species.
Inermonephrys Fauchald 1968a, Nephtys (Aglaophamus) Invalid Genera
inermis Ehlers 1887; 3 species.
Aglaopheme Kinberg 1866b, see Aglaophamus
Eversible pharynx without papillae. Interramal cirri
Aonis Savigny 1822, see Nephtys
involute, acicula distally hooked.
Diplobranchus Quatrefages 1865, see Nephtys
Micronephthys Friedrich 1937, Nephihys minuta Theel Pellucidaria Sveshnikov 1959, larval forms, no species
1 879; 5 species. named.
Eversible pharynx with 14 rows of papillae. Inter- Portelia Quatrefages 1865, see Nephrys
ramal cirri reduced or absent; acicula blunt-tipped,
but not capped. FAMILY SPHAERODORIDAE MALMGREN 1867
Nephtys Cuvier in Audouin and Milne Edwards 1833b, Small Phyllodocida with short and thick or long,
N. hombergii Savigny 1818; 50 species. relatively slender bodies. Two to six antennae and one
Eversible pharynx with 22 rows of papillae. Inter- pair of tentacular cirri present. Eversible pharynx
ramal cirri recurved; acicula in most forms with a unarmed. Uniramous parapodia with simple or com-
distal cap. posite setae. Dorsum with two to many rows of l arge
spherical tubercles.
The sphaerodorids have been overlooked in most
Taxonomic Notes
collections. They turn out to be quite frequent in deep-
Identification of nephtyids to genus can be tricky water samples, which tend to be better treated than
on small specimens; these tend to have small, nearly shallow-water ones. It is probably only a question of
straight interramal cirri and one could identify these time before they are found in relatively large numbers
as Micronephthys, and the corresponding adults as also in shallow water samples. Characteristically, they
Nephtys. Identification to species is not difficult, but are short and grub-shaped or more slender, vermiform,
care must be taken that appropriate segments are com- and cannot be confused with any other group of poly-
pared with each other (or with illustrations), since the chaetes, except perhaps Sphaerosyllis (Syllidae), which
shape of the parapodia changes along the length of the has similar rows of tubercles, in the latter case, the
body. To overcome the problem of varying body-length, dorsal cirri, along the dorsum. The setal structures,
comparisons should be made on distinct fractions of however, are quite different. Fauchald (1974b) re-
the bodies (i.e. first third, second third and last third). viewed the group.
Key to Genera
Ia. Both dorsal and ventral surfaces smooth Levidorum
lb. At least two rows of dorsal macrotubercles 2
2a (I b). Macrotubercles with terminal papillae 3
2b (lb). Macrotubercles distally rounded 6
3a (2a). Macrotubercles in four rows; terminal papillae very short Sphaerephesia
3b (2a). Macrotubercles in two rows; terminal papillae long 4

4a (3b). All setae simple Sphaerodorum

4b (3b). At least some setae composite 5
5a (4b). All setae composite apart from the recurved hooks in the first setiger Ephesiella
5b (4b). Both composite and simple setae in all setigers; apart from the first, which has simple recurved
hooks Ephesiopsis
6a (2b). Macrombercles stalked 7
6b (2b). Macrotubercles sessile 8
7a (6a). Median antenna as long as, or longer than, the lateral antennae Clavodorum
7b (6a). Median antenna shorter than the lateral antennae Sphaerodoridium
8a (6b). All setae composite Sphaerodoropsis
8b (6b). All setae simple Commensodorum
Generic Definitions Sphaerodoridium Lbtzen 1961, Sphaerodorum cla-

paredii Greeff 1866; only species.
Clavodorum Hartman and Fauchald 1971, C. at-
Stalked macrotubercles without terminal papillae.
lanticum Hartman and Fauchald 1971; 5 species.
Anterior end with a short median and two pairs of
Stalked macrotubercles in six or eight rows; macro-
longer lateral antennae. Setae composite.
tubercles without terminal papillae. Anterior end with
long median antenna and two pairs of lateral antennae. Sphaerodoropsis Hartman and Fauchald 1971, Sphaer-
Setae composite. odorum sphaerulifer Moore 1911; 20 species.
Four or more rows of macrotubercles without ter-
Commensodorum Fauchald 1974b, Sphaerodoridium minal papillae. Anterior end with a median and two or
commensalis Lutzen 1961; only species.
three pairs of lateral antennae. Setae composite.
Sessile macrotubercles in four rows; macrotubercles
without terminal papillae. Anterior end with a median Sphaerodorum Orsted 1843b, Ephesia gracilis Rathke
and two pairs of lateral antennae; all anterior appen- 1843; 5 species.
dages short. Setae simple. Four rows of sessile macrotubercles with terminal
papillae. Anterior end with a median and two pairs of
Ephesiella Chamberlin 1919c, Sphaerodorum abyss- lateral antennae. Setae simple, including in most species
arum Hansen 1878; 9 species. large recurved hooks in the first setiger.
Two rows of macro- and two rows of microtubercles
on the dorsum; the sessile macrotubercles with terminal Invalid Genera
papillae. Anterior end with a median and one or two
Bebryce Johnston 1844, see Sphaerodorum
pairs of lateral antennae. A large recurved hook present
Ephesia Rathke 1843, see Sphaerodorum
in the first setiger in most species; all other setae
Hypephesia Perrier 1897, see Ephesiella
composite.
Pollicita Johnston 1845, see Sphaerodorum
Ephesiopsis Hartman and Fauchald 1971, E. guayanae Thysanoplea Schmidt 1857, questionably Sphaerodorum
Hartman and Fauchald 1971; only species.
Two rows of macro- and two rows of microtubercles FAMILY TOMOPTERIDAB GRUBE 1848
on the dorsum; the sessile macrotubercles with terminal Pelagic, transparent, flattened Phyllodocida. Pros-
papillae. Anterior end with a median and two pairs of tomium fused with the first two segments; two antennae.
lateral antennae. A large recurved hook in the first Eversible pharynx unarmed. First segment well de-
setiger; setae in other setigers both simple and com- veloped in juveniles and reduced in adults. Second
posite. segment in adults with a pair of long tentacular cirri
supported by long acicula; setae absent. Parapodia
Levidorum Hartman 1967, L. scotiarum Hartman 1967;
biramous with elongated bases and short, foliose rami.
only species.
Glands of different kinds in the parapodia.
All tubercles and papillae absent. Anterior end blunt;
The tomopterids have been referred to the phyllo-
anterior appendages absent. Parapodia with two or three
dociform group of polychaetes, but differs rather sharply
acicula; setae semicomposite or simple.
from these in the structure of the anterior end (Uschakov
Sphaerephesia Fauchald 1972, S. longiseta Fauchald 1972) and appears best considered as a free-standing
1972; 3 species. family in the order Phyllodocia. A number of genera
Four rows of sessile macrotubercles with short have been described, but it appears best to consider
terminal papillae. Anterior end with a median and the family as consisting of only two genera. Recent
two or three pairs of lateral antennae. Setae com- revisions include Dales and Peter (1972) and Uschakov
posite. (1972).

Key to Genera
Ia. Tentacular cirri of second segment much longer than the body; parapodial fins restricted to the
far distal part of each ramus Enapteris
l b. Tentacular cirri as long as, or barely longer than the body; parapodial fins surround the distal part
of the rami Tomopteris
Generic Definitions
Enapteris Rosa 1908, Tomopteris euchaeta Chun 1887;

only species.
Parapodial rami with foliose lobes along the distal
margin only. Tentacular cirri much longer than body.
Tomopteris Eschscholtz 1825, T. onisciformis Esch-

Schultz 1825; 40 species.
Parapodia rami with foliose lobes surrounding most
of the rami. Tentacular cirri shorter than, or about
as long as the body.
Invalid Genera
Briaraea Quoy and Gaimard 1827, questionably

Tomopteris
Escholtzia Quatrefages 1865, see Tomopteris
Johnstonella Gosse 1853, see Tomopteris
FAMILY TYPHLOSCOLECIDAE ULIANIN 1878
Pelagic, transparent, fusiform or torpedo-shaped

Phyllodocida. Prostomium without appendages; large
foliose nuchal lobes present. Two pairs of tentacular
cirri. Eversible pharynx unarmed, but with retort-
organ. Parapodia uniramous, with large foliose dorsal
and ventral cirri; setae simple.
The typhloscolecids are poorly studied in terms of
FIGURE 27. (A), Family TYPHLOSCOLECIDAE, Travisiopsis
their biology and even basic biological information is lobifera, after Uschakov, 1972, about 12.5x; (B), Travisiopsis
missing. Recent reviews are given by Uschakov (1972) levinseni, anterior end, cirri on left side removed, after Uscha-
and Dales and Peter (1972). kov, 1972, 25x.
Key to Genera
la. Prostomium with ciliated ridges dorsally and ventrally Typhloscolex

lb. Prostomium without ciliated ridges 2
2a (lb). Median prostomial papilla on the dorsal side present Travisiopsis
2b (Ib). Prostomium without median dorsal papilla Sagitella
Generic Definitions
Sagitella Wagner 1872, S. kowalewskii Wagner 1872; Prostomium without ciliated ridges; median antenna
only species. present (may be indistinct).
Prostomium without ciliated ridges; median antenna
Typhloscolex Busch 1851, T. muelleri Busch 1851;
absent.
6 species.
Travisiopsis Levinsen 1885, T. lobifera Levinsen 1885; Prostomium with dorsal and ventral ciliated ridges;
6 species. median antenna present.

Taxonomic Notes the overall external similarity to the short-bodied

amphinomids, found the differences sufficient to erect
The above definitions are after Uschakov (1972:
a new family for the single genus. Manton (1967:10
220-225); there appears to be differences in the struc-
and 21) remarked upon the unique construction of the
ture of the nuchal organs in this family, and this may
eversible pharynx in Spinther compared to all other
be of value as a generic character in a future revision.
polychaetes. The genus also differs sharply from the
order Amphinomida in general and the erection of a
Invalid Genera separate order to contain this family seems warranted.
The new order, Spintherida, is listed below (p. 103).
Acicularia Langerhans 1878, see Sagitella
Nuchubranchiata Treadwell 1928, see Travisiopsis
FAMILY AMPHINOMIDAE SAVIGNY 1818
Plotobia Chamberlin 1919c, see Travisiopsis
Amphinomida with either elongate or ovate and
flattened bodies. One to five antennae present; palps
ORDER AMPHINOMIDA
present. Noto- and neurosetae in tufts, notosetae pro-
Prostomium distinct, a caruncle present and at least tective spinous setae, at least in part. Branchiae in
one antenna. Pharynx with a muscular, rasplike ever- branching tufts.
sible ventral pad; jaws absent. Parapodia distinct with The amphinomids are common in shallow water.
branching branchiae on at least some setigers. They generally are referred to as fire-worms, since
The family Spintheridae usually is associated with the spines can lead to general discomfort and infections,
the Amphinomidae (Fauvel 1923a; Hartmann-Schroder if they break off within the inflicted cuts, The amphi-
1971). The external resemblance of the ectoparasitic nomids generally are highly colored shallow-water
spintherids to the euphrosinids first were remarked animals, with very distinct color patterns (cf. Fauvel
upon by Sars (1850:210) and next by Johnston (1865: 1953), but are also present, in less showy editions,
127-128). Augener (1913a:87), while recognizing in deeper water.
Key to Genera
Ia. Caruncle completely absent; neurosetae simple hooks Hipponoa
l b. Caruncle present, usually well developed (may be difficult to discern in some species); neurosetae
otherwise 2
2a (lb). Body ovate or fusiform 3
2b (I b). Body elongated with parallel sides and usually abruptly tapering anteriorly and posteriorly 10
3a (2a). Branchiae present on all setigers Branchamphinome
3b (2a). Some setigers (anterior or posterior) without branchiae 4
4a (3b). One dorsal citrus per notopodium 5
4b (3b). Two dorsal cirri per notopodium 7
5a (4a). Caruncle with three parallel longitudinal ridges Benthoscolex
5b (4a). Caruncle long and folded with indistinct lateral folds 6
6a (5b). Eyes absent, first pair of branchiae larger than the following ones Bathychloeia
6b (5b). Eyes present, first branchiae not larger than the following ones Chloeia
7a (4b). Caruncle with high central ridge and two wide flattened lateral folds Notopygos
7b (4b). Lateral lobes of caruncle small and hidden under the central ridge, or absent 8
8a (7b). Caruncle wedge-shaped, without crest and folds Sangiria
8b (7b). Caruncle with crest and folds 9
9a (8b). Caruncle high, loosely plaited and rugose Chloenopsis
9b (8b). Caruncle low, narrowly plaited with a crenulated plate Parachloeia
l0a (2b). Caruncle small and inconspicuous, stretching through maximally three segments 11
l0b (2b). Caruncle large and conspicuous, stretching through at least three segments 14
1 Ia (l0a). Branchiae present on all segments from the second or third 12
l lb (l0a). Branchiae limited to the anterior part of the body 13
12a (11a). Caruncle broadly triangular or cordate Amphinome
12b (11a). Caruncle narrow and elongated Pareurythoe
l 3a (I lb. First segment with large, anteriorly directed hooks Paramphinome
13b (lib). First segment without hooks Linopherus
FIGuRE 28. (A), Family AMPHINOMIDAE, Eurythoe complanata, San Felipe, Golfo de California, intertidal, lOx; (B), median
parapodium of the above, lOx; (C), Family EUPHROSINIDAE, Euphrosine borealis, Murchinson Sound, Greenland, 100 m,
lOx; (D), Euphrosine sp., off Point Loma, California, median parapodium, 19x; (E), Family SPINTHERIAE, Spinther sp.,
Point Barrow, Alaska, dredged, Sx.

14a (l0b). Caruncle longer than wide with a large smooth, sinuous median ridge nearly covering the narrow
lateral parts Eurythoe
14b (l0b). Caruncle about as long as wide or wider, median ridge, if present, narrow 15
I5a (14b). Caruncle without distinct median ridge, with a few deep transverse folds Hermodice
15b (14b). Caruncle with a distinct, smooth narrow ridge 16
16a (15b). Furcate setae absent Pherecardia
I6b (15b). Furcate setae present Pherecardites
Generic Definitions folds. Dendritically branched branchiae. Dorsal cirri

single, eyes present.
Amphinome Brugui8re 1789, Aphrodita rostrata Pallas
1776; 12 species. Hermodice Kinberg 1857, Aphrodita carunculata
Body long; caruncle limited to the two-three first Pallas 1766; 4 species.
segments, broadly triangular or cordate. Dendritically Body long; caruncle about as wide as long, without
branched branchiae on all segments from the second a distinct median ridge, with several transverse folds.
or third. Branchiae dendritically branched, bushy in appearance.
Bathychloeia Horst 1910, B. sibogae Horst 1910; only Dorsal cirri single, eyes present.
species. Hipponoa Audouin and Milne Edwards 1830, H.
Body short and ovate; caruncle large, folded with gaudichaudi Audouin and Milne Edwards 1830;
l ateral folds. Bipinnate branchiae from segment 5, 3 species.
first pair larger than the others. Dorsal cirri single. Body short and ovate; caruncle absent. Branchiae
Eyes absent. dendritically branched, bushy. Dorsal cirri single,
Benthoscolex Horst 1912, B. coeca Horst 1912; 2 eyes present. Notosetae large, retractile curved hooks.
species.
Linopherus Quatrefages 1865, Amphinome incarun-
Body short and ovate; caruncle with three longitu- culata Peters 1854; 14 species.
dinal parallel ridges. Dendritically branched branchiae
Small species, but elongated bodies in most forms;
from setiger 6, especially strongly developed on poste-
caruncle small and inconspicuous, reported absent in
rior setigers. Dorsal cirri single, eyes absent.
some forms. Branchiae tufted, present on some anterior
Branchamphinome Hartman 1967, B. antarctica Hart- setigers only. Dorsal cirri single, eyes present or
man 1967; only species. absent.
Body ovate; caruncle with three digitate posteriorly
Notopygos Grube 1855, N. crinita Grube 1855; 20
directed lobes. Dendritically branched branchiae present
species.
on all setigers. Dorsal cirri single, eyes present.
Body ovate; caruncle with high central ridge and
Chloeia Savigny 1818, Aphrodita flava Pallas 1766; l arge flattened lateral lobes. Branchiae dendritically
19 species. branched. Dorsal cirri double; eyes present.
Body ovate, caruncle large, with indistinct lateral
folds, much longer than wide. Branchiae pennate. Parachloeia Horst 1912, P. marmorata Horst 1912;
Dorsal cirri single, eyes present. only species.
Body ovate; caruncle low, narrowly plaited with
Chloenopsis new name, Chloenea atlantica McIntosh crenulated plates. Branchiae poorly developed, with
1885; only species. a few filaments only. Dorsal cirri double, eyes present.
Body ovate, caruncle high, loosely plaited and rugose.
Branchiae pennate. Dorsal cirri double; eyes present. Paramphinome Sars 1869, Hipponoe jeffreysii Mc-
Chloenea Kinberg, 1867c is considered a synonym Intosh 1868; 6 species.
of Chloeia. Chloenea McIntosh, 1885 differs gen- Small forms, but long-bodied; caruncle short,
erically from Kinberg's genus and needs a new name. Y-shaped or elongated. Branchiae tufted, limited to
Chloenopsis is here proposed as the generic name for anterior setigers. Dorsal cirri single; eyes present or
the species originally named Chloenea atlantica McIn- absent. First setiger with anteriorly directed hooks.
tosh, 1885.
Pareurythoe Gustafson 1930, P. japonica Gustafson
Eurythoe Kinberg 1857, E. capensis Kinberg 1857; 1930; 9 species.
10 species. Body long; caruncle small, elongated, sinuous.
Body long; caruncle long with a thick, sinuous, Branchiae dendritically branched, present on most
smooth median ridge covering the narrow folded lateral of the body. Dorsal cirri single, eyes present.

Pherecardia Horst 1886, Hermodice striata Kinberg Didymobranchus Schmarda 1861, indeterminable
1857; 4 species. Eucarunculatus Malaquin and Dehorne 1907, see
Body long; caruncle with a narrow smooth ridge Pherecardia
bordered by wide lateral lobes with deep parallel folds Lenora Grube 1878, see Amphinome
on both sides. Branchiae bushy. Dorsal cirri single; Lirione Kinberg 1867c, see Notopygos
eyes present. Lycaretus Kinberg 1867c, see Eurythoe
Metamphinome Treadwell 1940, see Hipponoa
Pherecardites Horst 1912, P. parva Horst 1912; 2
Pleione Savigny 1818, possibly Amphinome
species.
Pseudeurythoe Fauvel 1932, see Linopherus
Body long; caruncle with median axis and lateral
Rostraria Haecker 1898, larval forms
lamellae directed posteriorly. Bushy branchiae present
Strategis Kinberg 1867c, see Chloeia
from the first setiger. Dorsal cirri single, eyes present.
Thesmia Kinberg 1867c, see Chloeia
Furcate neurosetae present.
Thetisella Baird 1870, larval forms
Sangiria Horst 1911, S. hystrix Horst 1911; only Veleda Castelnau 1842, indeterminable
species. Zothea Risso 1826, indeterminable
Body ovate; caruncle wedge-shaped without crest
and folds. Branchiae with few filaments. Dorsal cirri
FAMILY EUPHROSINIDAE WILLIAMS 1851
double; eyes absent.
Amphinomida with short and thick bodies.One pair
of antennae; palps absent. Neurosetae in tufts, notosetae
Invalid Genera
in transverse rows on the dorsum; branching branchiae
Amphibranchus Kinberg 1867c, see Hermodice in rows between the notosetae.
Asloegia Kinberg 1867c, see Amphinome Euphrosinids often are considered in the amphi-
Blenda Kinberg 1867c, see Eurythoe nomids SENSU LATU. The two families are closely
Chloenea Kinberg 1867c, see Chloeia related, but the euphrosinids make up a distinct, com-
Chloochaeta Kinberg 1867c, see Chloeia pact group of forms, and it appears best to treat them
Colonianella Kinberg 1867c, see Amphinome separately.
Key to Genera
Notosetae bifurcate, with cylindrical shafts Euphrosine
Notosetae flattened, smooth paleae Palmyreuphrosyne
Generic Definitions smooth and cylindrical, which would ally the genus
closely to Palmyra rather than to the amphinomidlike
Euphrosine Savigny 1818, E. myrtosa Savigny 1818;
forms with their ventral plate-muscle pharynx. As
40 species.
indicated by Orrhage on several occasions, structure
Short-bodied forms with short prostomium, caruncle
of the pharynx may not be an overwhelmingly strong
with three longitudinal lobes. Branchiae in transverse
character, and it appears best to await further study
rows in the dorsum of each segment. Setae include
of these forms to decide the question. In the mean-
capillaries, and furcate setae.
ti me, Palmyreuphrosyne is maintained where it was
Palmyreuphrosyne Fauvel 1913, P. paradoxa Fauvel placed by its original describer, Fauvel (1913).
1913; 2 species.
Short-bodied forms with an elongated caruncle. Invalid Genus
Pectinate branchiae in three dorsal groups per segment.
Lophonota Costa 1841, see Euphrosine
Parapodia transverse ridges. Notosetae flattened smooth
paleae.
ORDER SPINTHERIDA
Body ovate. Prostomium with a median antenna.

Taxonomic Note Pharynx retractable and cylindrical (resembles a tur-
Palmyreuphrosyne appears, as noted by Fauvel bellarian pharynx). Notopodia represented by mem-
(1913) to combine characters of the euphrosinids with branous ridges supported by simple or furcate setae.
those of the palmyrids. Fauvel indicates that the ever- Neuropodia with composite, strongly curved hooks.
sible pharynx of Palmyreuphrosyne should be ventral, Ectoparasitic on sponges.

Family Spintheridae
As the order; the family is known for a single
genus, Spinther Johnston 1845, with the genotype,
S. oniscoides Johnston 1845. About 12 species pres-
ently are recognized.
Invalid Genera
Cryptonota Stimpson 1854, see Spinther

Oniscosoma Sars 1850, see Spinther
ORDER EUNICIDA
Prostomium distinct, with or without appendages.

Eversible pharynx ventrolateral, strongly muscular
and with at least one pair of jaws. Parapodia distinct,
with strongly developed neuropodia and reduced
notopodia.
Superfamily Eunicea
Two to five pairs of lateral jaws (maxillae) and
usually one pair of lower jaws (mandibles).
FAMILY ONUPHIDAE KINBERG 1865
Eunicea with two frontal and five occipital antennae.

Maxillary carriers short; third carrier absent. Maxilla I
smooth and curved. Notopodia represented by the base
of the branchiae and the dorsal cirri, often supported
by internal acicula. Setae include composite and pseudo-
composite hooks and spinigers, pectinate setae, limbate
setae and subacicular hooks.
Most onuphids are tubicolous, some of them carry
the tube around (Hyalinoecia); others are sessile, but
may be able to leave their tubes in emergencies (Schafer
1972). All species appear to be scavenging and feed
on both plant and animal debris floating past their tube
openings; others may actively hunt for debris. Onuphids
tend to be common at all depths, and are, next to the
FIGURE 29. (A), Family ONUPHIDAE, Nothria elegans, off
lumbrinerids, the family of Eunicea best represented Santa Barbara, California, 20 m, 15x; (B), maxillae of the
in deep water. Recent revisions were made by Fauchald above, 25x; (C), third parapodium of the above, 35x; (D),
(1968b, 1972). median parapodium of the above, 25x.
Key to Genera
Ia. Two or more anterior setigers with prolonged parapodia and modified setae 2
lb. Anterior parapodia not prolonged, or only first parapodium longer than the following ones 4
2a (la). Two modified anterior setigers; these with uni- or bidentate hooks and capillary setae . . Paranorthia
2b (Ia). More than two modified anterior setigers 3
3a (2b). Modified parapodia with strongly curved, grapple-hook shaped setae Rhamphobrachium
3b (2b). Modified parapodia with composite falcigers Americonuphis
4a (III). Branchiae in part spiralled 5
4b (lb). Branchiae pectinate, simple or absent 7
5a (4a). Tentacular cirri absent Epidiopatra
5b (4a). Tentacular cirri present 6
6a (5b). Frontal antennae short and conical, all dorsal cirri digitate Diopatra

6b (5b). Frontal antennae long and slender, at least some dorsal cirri foliose Heptaceras
7a (4b). Tentacular cirri absent 8
7b (4b). Tentacular cirri present 9
8a (7a). Branchiae present Hyalinoecia
8b (7a). Branchiae absent Paronuphis
9a (7b). Branchiae in part pectinate Onuphis
9b (7b). Branchiae simple or absent 10
l0a (9b). Branchiae absent, dorsal cirri foliose in some anterior setigers Paradiopatra
l0b (9b). Branchiae present or absent, all dorsal cirri digitate Nothria
Generic Definitions Paradiopatra Ehlers 1887, P. fragosa Ehlers 1887;

2 species.
Americonuphis Fauchald 1973, Diopatra magna An-
Frontal antennae short and conical, tentacular cirri
drews 1891; 2 species.
present. Anterior setigers and setae not modified. Bran-
Frontal antennae short and conical; tentacular cirri
chiae absent, dorsal cirri foliose in some setigers.
present. Five or more anterior setigers modified with
composite falcigers. Branchiae pectinate; dorsal cirri Paranorthia Moore 1903, P. brevicornuta Moore 1903;
digitate. 5 species.
Diopatra Audouin and Milne Edwards 1833b, D.
present. Two modified anterior setigers present with
amboinensis Audouin and Milne Edwards 1833b; 40
uni- and bidentate hooks and capillary setae. Branchiae
species.
simple and straplike or bifid, dorsal cirri digitate.
present. Anterior setigers and setae not modified. Paronuphis Ehlers 1887, P. gracilis Ehlers 1887;
Branchiae spiralled, dorsal cirri digitate. 5 species.
Epidiopatra Augener 1918, E. hupferiana Augener
absent. Anterior setigers and setae not modified.
1918; 5 species.
Branchiae absent, dorsal cirri digitate.
absent. Anterior setigers and setae not modified, Rhamphobrachium Ehlers 1887, R. agassizi Ehlers
branchiae spiralled, dorsal cirri digitate. 1887; 10 species.
Heptaceras Ehlers 1868, Diopatra phyllocirrus Schmarda
present. Two or more anterior setigers modified with
1861; only species.
strongly curved, grapple-hooklike setae. Branchiae
Frontal antennae long and slender; tentacular cirri
pectinate, dorsal cirri digitate.
present. Anterior setigers and setae not modified.
Branchiae spiralled, dorsal cirri foliose in some setigers.
Taxonomic Notes
Hyalinoecia Malmgren 1867, Nereis tubicola O.F.
Muller 1788; 20 species. Nothria often is considered a synonym of Onuphis.
Frontal antennae short and conical; tentacular cirri This is probably correct; species of both genera re-
absent. Anterior setigers and setae not modified. semble each other closely. The separation is retained
Branchiae present, simple and straplike in most species; here for practical reasons, a final decision of this ques-
dorsal cirri digitate. tion will have to await the study of larger materials
than is presently available.
Nothria Malmgren 1867, Onuphis conchylega Sars
1835; 40 species.
Frontal antennae short and conical; tentacular cirri Invalid Genera
present. Anterior setigers and setae not modified.
Leptoecia Chamberlin 1919c, see Paronuphis
Branchiae simple and straplike or absent, dorsal cirri
Nereitube Blainville 1828, see Hyalinoecia
digitate. Northia Johnston 1865, see Nothria
Onuphis Audouin and Milne Edwards 1833b, O. Trapodia Baird 1870, see Onuphis
eremita Audouin and Milne Edwards 1833b; 60
species.
FAMILY EUNICIDAE SAVIGNY 1818
Frontal antennae short and concial, tentacular cirri
present. Anterior setigers and setae not modified. Eunicea with from one to five occipital antennae.
Branchiae pectinate, dorsal cirri digitate. Maxillary carriers short; third carrier absent. Maxilla I

smooth and curved. Notopodia represented by branchiae substrates and thus with shallow water (Fauchald 1969,
and dorsal cirri, sometimes supported by internal 1970). Generally, the eunicids are considered carni-
acicula. Setae include composite falcigers and spinigers, vores, but some may be scavengers or live on large
li mbate setae, pectinate setae and subacicular hooks. detrital particles. Tube-building is known for some
The eunicids are among the largest of polychaetes, species; others are burrowing into limestone or other
some Eunice aphroditois have been reported as long calcium carbonate substrates. Major revisions include
as two meters. Most species are associated with hard Hartman (1944a) and Fauchald (1970).
Key to Genera
Ia. Five occipital antennae present 2
lb. One to three occipital antennae present 6
2a (Ia). Tentacular cirri present 3
2b (la). Tentacular cirri absent 5
3a (2a). Subacicular hooks absent Palola
3b (2a). Subacicular hooks present 4
4a (3b). Composite setae falcigers Eunice
4b (3b). Composite setae spinigers Euniphysa
5a (2b). Branchiae present Marphysa
5b (2b). Branchiae absent Paramarphysa
6a (1 b). One occipital antenna present Nematonereis
6b (1 b). Three occipital antennae present Lysidice
Generic Definitions
Eunice Cuvier 1817 (NOMEN CONSERVANDUM) NereiS
aphroditois Pallas 1788; 170 species.
Five occipital antennae; tentacular cirri present;
branchiae present. Setae include limbate setae, pectinate
setae, composite falcigers and subacicular hooks.
Euniphysa Wesenberg-Lund 1949, E. aculeata Wesen-
berg-Lund 1949; only species.
Five occipital antennae; tentacular cirri present;
setae, composite spinigers and subacicular hooks.
Lysidice Savigny 1818, L. ninetta Audouin and Milne

Edwards 1833a; 10 species.
Three occipital antennae; tentacular cirri and branchiae
absent. Setae include limbate setae, pectinate setae,
composite falcigers and subacicular hooks.
Marphysa Quatrefages 1865, Nereis sanguinea Mon-
tagu 1815; 50 species.
Five occipital antennae; tentacular cirri absent,
setae, composite spinigers and falcigers and subacicular
hooks.
Nematonereis Schmarda 1861, N. unicorns Schmarda

1861; 2 species.
One occipital antenna; tentacular cirri and branchiae
absent. Setae include limbate and pectinate setae, com-
posite spinigers and subacicular hooks.
FIGURE 30. (A),Family EUNICIDAE, Eunice antennata, Point
Palola Gray 1847, Eunice siciliensis Grube 1840; Loma, California, intertidal, tOx; (B), subacicular hook of the
4 species. above, 100x; (C), median parapodium of the above, 25x.

Five occipital antennae; tentacular cirri and branchiae Nauphanta Kinberg 1865, see Marphysa
present. Setae include limbate and pectinate setae and Nausicaa Kinberg 1865, see Marphysa
composite falcigers. Nereidice Blainville 1828, see Lysidice
Nereidonta Blainville 1828, see Eunice, Marphysa
Paramarphysa Ehlers 1887, P. longula Ehlers 1887; and Palola
4 species. Palpiglossus Wagner 1885, indeterminable
Five occipital antennae; tentacular cirri and branchiae Pseudopalolo Friedlander in Woodworth 1907, see
absent. Setae include limbate and pectinate setae, Lysidice
composite falcigers and subacicular hooks. Tibiana Lamarck 1816, see Eunice
Taxonomic Notes FAMILY LUMBRINERIDAE MALMGREN 1867

Palola frequently is considered a subgenus of, or Eunicea without prostomial appendages, but some-
synonymous with Eunice. The latter is a large genus, times with one to three nuchal papillae emerging from
and anything that can be done to subdivide it, appears a pocket between the pro- and peristomium. Maxillary
to be of value. Members of Palola differ consistently carriers short, third carrier absent. Maxilla I smooth
and uniquely from the rest of the family not only in and curved. Notopodia absent or represented by small,
the characters listed above, but also in the deep scoop- button-shaped projections, sometimes with internal
shape of the mandibles. It thus appears valuable to acicula (but see Kuwaita below). Setae include limbate
retain the distinction, not least since it appears related setae, simple and composite hooks. Subacicular hooks
to the ecology of the contained species. and pectinate setae are absent.
Paramarphysa and Marphysa are very similar and Most lumbrinerids are free-living, burrowing forms
can be separated only as indicated in the key; the in sand or mud or between algal hold-fasts and plant-
separation is maintained here provisionally. roots. They are among the most common polychaetes
in sandy and muddy bottoms in shelf-depths, but also
occur in numbers in deep-water areas. Identification
Invalid Genera of lumbrinerids is dependent on very accurate work
and is not altogether easy. Most dependable appears
Amphelothrix Chamberlin 1919c, error for Aphelothrix, to be the structure of the anterior setigers, the structure
see Marphysa of the jaw-apparatus and the detailed structure of the
Amphiro Kinberg 1865, see Marphysa hooks. Length and shape of the parapodial lobes also
Amphiron Chamberlin 1919c, error for Amphiro, are valuable characters, but care must be taken in
see Marphysa noting the exact location of the parapodium examined
Aphelothrix Chamberlin 1919c, see Marphysa in relation to the total length of the animal. This often
Blainvillea Quatrefages 1865, see Nematonereis is difficult since the lumbrinerids fragment readily.
Eriphyle Kinberg 1865, see Eunice An important character in most keys to species concerns
Eunice Rafinesque 1815, NoMEN NuDuM (Eunice Cuvier the length of the posterior parapodial lobes in relation
1817 is the valid name for the main genus in this to the length of the anterior ones; usually stated as
"prolonged posterior lobes." The key feature in this
family, fide Fauvel 1918:338).
Heteromarphysa Verrill 1900, indeterminable character, is that the posterior lobes have to be dis-
Leodice Savigny 1818, see Eunice tinctly longer than the anterior ones. With a fragmented
Lithognatha Stewart 1811, see Palola specimen this may be impossible to observe; in such
Lycidice Williams 1851, error for Lysidice cases, it is of the utmost importance that both paths
Macdufa McIntosh 1885, see Marphysa in a key be explored; in other words, that no assumption
Mayeria Verrill 1900, see Eunice is made about the structure of the posterior lobes.
Key to Genera
la. Parapodia with distinct dorsal cirri Kuwaita
lb. Dorsal cirri absent 2
2a (lb). Pharyngeal apparatus absent Ophiuricola
2b (lb). Pharyngeal apparatus present 3
3a (2b). Branchiae absent Lumbrineris
3b (2b). Branchiae present Ninoe
H
FIGURE 31. (A), Family IPHITIMIDAE, Iphitime loxorhynchi, from Loxorhynchus grandis, Santa Catalina Island, California,
26x; (B), Family LUMBRINERIDAE, Lumbrineris californiensis, off Point Firmin, California, 30 m, anterior composite hooded
hook, 385x; (C), posterior simple hooks of the above, 385x; (D), anterior end of the above, 16x; (E), 100th parapodium of the
above, 53x; (F), third parapodium of the above, 53x; (G), Family ARABELLIDAE, Arabella iricolor, Dillon Beach, California,
intertidal, 16x; (H), fifth parapodium of the above, 95x; (1), Family LYSARETIDAE, Oenone fulgida, Bahia Magdalena, Baja
California, 30 m, 26x; (1), fifteenth parapodium of the above, 52x.

Key to Genera Fide Orensanz (1974a)
Ia. Hooded hooks distally bidentate 2

lb. Hooded hooks, if present, distally multidentate (anterior parapodia may have incompletely formed
hooks) 3
2a (Ia). Maxilla IV with a series of articulated spines; mandibles posteriorly bifid; maxillary carriers large
and posteriorly expanded Lumbrineriopsis
2b (Ia). Maxilla IV without denticles or articulated spines; mandibles posteriorly entire; maxillary carriers
robust, subtriangular and not expanded posteriorly Lumbrinerides
3a (I b). Digitiform branchiae emerging from the postsetal lobes on a number of anterior setigers; maxillae
IV, or III and IV usually with denticulated cutting edge Ninoe
3b (Ib). If branchial structures present, they are in a different position; maxillae III and IV either smooth
or with a few large teeth 4
4a (3b). Anterior parapodia with composite hooded hooks; maxilla II with three large teeth; maxilla IV
greatly expanded with a central thin area Augeneria
4b (3b). Parapodia with or without composite setae. Maxillae different Lumbrineris
Generic Definitions we are using to define the many species, so that we

may use knowledge of this variability when we attempt
Kuwaita Mohammad 1973, K. magna Mohammad
to describe more genera in this family.
1973; only species.
The genus Kuwaita is isolated in the family and
Lumbrinerids with three nuchal papillae; dorsal cirri
may belong to a different, perhaps undescribed family.
present and elongate in posterior segments; branchiae
absent. Jaw apparatus present.
Lumbrineris Blainville 1828, L. latreilli Audouin and Invalid Genera

Milne Edwards 1834; 158 species. Aotearia Benham 1927, see Lumbrineris
Lumbrinerids without or with one to three nuchal Augeneria Monro 1930, see Lumbrineris (but see
papillae; dorsal cirri and branchiae absent. Jaw ap- above)
paratus present. Cenogenus Chamberlin 1919c, see Lumbrineris
Ninoe Kinberg 1865, N. chilensis Kinberg 1865; 28 Eranno Kinberg 1865, see Lumbrineris
species. Lumbriconereis Grube 1840, see Lumbrineris
Lumbrinerids without nuchal papillae, dorsal cirri Scoletoma Blainville 1828, see Lumbrineris
absent; branchiae present. law apparatus present. Unciniseta Bidenkap 1907, see Lumbrineris
Zygolobus Grube 1863, see Lumbrineris
Ophiuricola Ludwig 1905, O. cynips Ludwig 1905; Zygophyllus Grube 1863, error for Zygolobus, see
only species. Lumbrineris
Lumbrinerids without nuchal papillae, dorsal cirri
or jaw apparatus.
FAMILY IPHITIMIDAE FAUCHALD 1970
Taxonomic Notes
Eunicea with a single pair of frontal antennae. One
Orensanz (1974a) described the two genera Lum- pair of maxillary carriers fused to maxilla I, which is
brineriopsis and Lumbrinerides with L. mucronata smooth and curved; third carrier is absent. Notopodia
(Ehlers 1908) and L. gesae Orensanz 1974a as type represented by simple or branched branchiae. Setae
species respectively. Both genera may be valid. The include simple and composite falcigers.
change in concept of the previously described Augeneria A single genus: Iphitime Marenzeller 1902, with
and Ninoe indicated in the alternative key above appears genotype I. doederleini Marenzeller 1902, and five
unfortunate and is not recommended for future adop- species are known.
tion. Admittedly the genus Lumbrineris as presently All Iphitimidae are inquilines in branchial chambers
accepted, contains a large number of rather different of crustaceans; their feeding modes are unknown; it
forms, but experience has taught me that surveys based is possible that they parasitize the crustacean host; but
on descriptions from the literature are considerably little damage seems to be present on the branchial
less valuable than ones based on actual specimens. tissue of the host. Pilger (1972) and Fauchald (1970)
It is now time to study the variability of the characters reviewed the family.

110
lb.
2a (la).
2b (la).
3a (2b).
3b (2b).
4a (lb).
4b (lb).
5a (4b).
5b (4b).
6a (5b).
6b (5b).
7a (6b).
7b (6b).
20 species.
i
NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY
Invalid Genera
Coelobranchus Izuka 1912, see Iphitime
Enonella Stimpson 1854, indeterminable
FAMILY ARABELLIDAE HARTMAN
Acicular spines absent

1944a
Eunicea without prostomial appendages. Maxillary
carriers long and narrow, a third carrier present. Maxilla
I smooth or basally dentate; notopodia absent. Setae
include limbate setae and in a number of cases, thick,
emergent spines.
Ia. Acicular spines present
Maxillary apparatus represented by a single rod

Four or five pairs of jaws present
Maxilla I distally falcate
Maxilla I distally dentate
Maxillary apparatus absent
Maxillary apparatus present, but often reduced
Five pairs of maxillae present
Two or three pairs of maxillae present
Mandibles fused and horseshoe-shaped
Mandibles two triangular plates
Maxillary carriers anteriorly bilobed and fused along most of their lengths
Maxillary carriers anteriorly rounded and fused along their whole length
Generic Definitions
Arabella Grube 1850, Nereis iricolor Montagu 1804;
Arabellids with five pa rs of maxillae and mandibles

present. Parapodia without acicular spines.
Biborin Chamberlin 1919a, B. ecbola Chamberlin
1919a; only species.
Arabellids without maxillary apparatus, mandibles
present. Parapodia without acicular spines.
Drilognathus Day 1960, D. capensis Day 1960; only
species.
Arabellids with the maxillary apparatus reduced to
a single rod. Parapodia with acicular spines.
Drilonereis Claparede 1870b, Lumbriconereis filum

Claparede 1868; 40 species.
Arabellids with four or five pairs of maxillae; mandi-
bles usually present. Parapodia with acicular spines;
maxilla I distally falcate.
Haematocleptes Wiren 1886, H. terebellides Wiren
1886; 2 species.
Arabellids with the maxillary apparatus consisting
of two pairs of small plates, mandibles present and
separate; unpaired carrier present. Acicular spines
absent.
Science Series 28
Arabellids resemble lumbrinerids closely in overall

body-construction, but differ sharply from the latter
in that they have long narrow maxillary carriers and
have three, rather than two carriers. Arabellids are
never tubicolous; they may be parasitic in other animals,
usually in other polychaetes, but also in echiurans. They
may be parasitic as juveniles and become free-living
as adults (Pettibone 1957c), or may stay parasitic
throughout life (Wiren 1886). Major revisions: Hartman
1944a; Pettibone 1957c; Fauchald 1970.
Key to Genera
2
4
Drilognathus
3
Drilonereis
Notocirrus
Biborin
5
Arabella
6
Oligognathus
Arabellids with two pairs of minute maxillae and

the carriers fused for part of their length. Acicular
spines absent.
Notocirrus Schmarda 1861, N. chilensis Schmarda
1861; 10 species.
Notopsilus Ehlers 1868, Lais acutus Kinberg 1865;

only species.
Arabellids with five pairs of maxillae and mandibles
present. Acicular spines absent. Maxillae I proximally
dentate.
Oligognathus Spengel 1882, O. bonelliae Spengel

1882; 2 species.
7
Labrorostratus
Haematocleptes
Labrorostratus Saint-Joseph 1888, L. parasiticus

Saint-Joseph 1888, only species.
Arabellids with four or five pairs of maxillae, mandi-

bles present. Maxillae I dentate to the tip. Acicular
spines present.
Arabellids with two or three pairs of maxillae present;

mandibles fused into horseshoe-shaped piece. Acicular
spines absent.
Taxonomic Notes
The generic sub-division is presently inconsistent
in that in the group of arabellids with emergent acicular
spines, forms with maxilla I distally falcate are con-

/976 THE POLYCHAETE WORMS III
sidered generically distinct from forms with maxilla I Lais Kinberg 1867, see Notopsilus
distally dentate. The corresponding separation should Laranda Kinberg 1865, indeterminable
split the genus Arabella so that the forms with maxilla I Maclovia Grube 1871 b, see Arabella
dentate should go to the genus Notopsilus, which has Pterothrix Chamberlin 1919c, see Notocirrus
been defined in the generic definitions, but not con-
sidered in the key. This genus has been largely disre- FAMILY LYSARETIDAE KINBERG 1865
garded; it is not clear whether the best procedure would Eunicea with one or three occipital antennae. Maxil-
be to revise Arabella to separate the two sets of forms,
l ary carriers long and narrow, a third carrier is present;
or to fuse the well-known genera Notocirrus and Dri-
maxilla I proximally smooth or dentate. Notopodia
lonereis to ensure generic conformity within the represented by large dorsal cirri supported by acicula.
family.
Setae include limbate setae and in some genera, bi-
dentate hooded setae.
Invalid Genera
Lysaretidae is a small family of rather large, mainly
Arabes Ehlers 1920, see Drilonereis tropical shallow-water polychaetes. The general ap-
Aracoda Schmarda 1861, see Arabella, Notocirrus pearance of the worms is that of a lumbrinerid, but the
and Lumbrineris large dorsal cirri are distinct, as is the jaw-apparatus.
Cenothrix Chamberlin 1919c, see Arabella Important revisions include Fauchald (1970) and Knox
Labidognathus Caullery 1914a, see Drilonereis and Green (1972).
Key to Genera
Ia. One distinct peristomial segment present Oenone

l b. Two distinct peristomial segments present 2
2a (lb). One short antenna present Tainokia
2b (Ib). Three antennae present 3
3a (2b). Proximal end of maxilla I dentate; distal end falcate Halla
3b (2b). Proximal end of maxilla I smooth or nearly smooth; each distal end with two large fangs . . Lysarete
Generic Definitions determined. Crossland (1924) investigated some of the

variability of the jaw-apparatus, but did not have large
Halla Costa 1844, Nereis parthenopeia delle Chiaje
enough materials to confirm or reject the presence of
1828; 3 species.
distinct sub-groups. Halla parthenopeia also has been
Lysaretids with two distinct peristomial rings, three
reported from cosmotropical areas and may be another
antennae and the distal end of maxilla I falcate. The
species-complex.
proximal end of maxilla I is dentate.
Lysarete Kinberg 1865, L. brasiliensis Kinberg 1865; Invalid Genera

2 species.
Lysaretids with two distinct peristomial rings, three Aenone Risso in Quatrefages 1865, indeterminable
antennae and the distal end of maxilla I divided into Aglaura Savigny 1818, see Oenone
two large fangs. Proximal end of maxilla I smooth. Aglaurides Ehlers 1868, see Oenone
Andromache Kinberg 1865, see Oenone
Oenone Savigny 1818, Aglaura fulgida Savigny 1818; Cirrobranchia Ehlers 1868, see Halla
4 species. Danymene Kinberg 1865, indeterminable
Lysaretids with one distinct peristomial ring, three Larymna Kinberg 1865, indeterminable
antennae and usually, distally falcate maxilla I. Prox- Plioceras Quatrefages 1865, see Halla
imal end of maxilla I dentate.
Tainokia Knox and Green 1972, T. iridescens Knox FAMILY DORVILLEIDAE CHAMBERLIN 1919C
and Green 1972; only species. Eunicea with two pairs of antennae. Maxillae consist
Lysaretids with two distinct peristomial rings, one
of one or two series of small jaw-pieces and paired
antenna and distally falcate maxilla I. Proximal end
carriers; mandibles present. Notopodia reduced, but
of maxilla I is dentate.
with setae and acicula in most forms. Setae include
simple and composite hooks, furcate and limbate setae.
Taxonomic Notes
Dorvilleids are mainly small polychaetes, considered
The number of species involved in the circumtropical most common in shallow water, but recently recovered
complex referred to as Oenone fulgida has not been i n increasing numbers from deeper water (Jumars 1974).

One pair of the antennae is referred to as palps; they The jaw-apparatus may also be reduced in some forms.
differ in structure from the other pair, in that they often Jumars (1974) has revised the generic classification
are articulated. Either one or both pairs may be reduced. of the family.
Key to Genera
Ia. Notacicula present 2
lb. Notacicula absent 3
2a (Ia). Furcate setae present Schistomeringos
2b (Ia). Furcate setaee absent Dorvillea
3a (lb). Furcate or geniculate setae present 4
3b (Ib). Furcate or geniculate setae absent 5
4a (3a). Palps well developed Protodorvillea
4b (3a). Second pair of antennae reduced or absent Meiodorvillea
5a (3b). Only simple acicular setae present Parophryotrocha
5b (3b). Both capillary and compound setae present 6
6a (5b). Setae of first setiger markedly different from the others Exallopus
6b (5b). Setae of first setiger similar to others 7
7a (6b). First antennae long and cirriform Apophryotrocha
7b (6b). First antennae reduced and papilliform Ophryotrocha
Generic Definitions Ophryotrocha Claparede and Mecznikow 1869, O.

puerilis Claparede and Mecznikow 1869; 11 species.
Apophryotrocha Jumars 1974, A. mutabiliseta Jumars Dorvilleids with antennae and palps reduced to
1974; only species. papilliform projections. Dorsal cirri without acicula,
Dorvilleids with very long, cirriform antennae and may be absent. Setae include capillaries and hetero-
well-developed palps. Dorsal cirri without acicula. gomph falcigers. Carriers and basal plates fused into
Setae of two kinds, capillaries and composite hetero- a characteristic ice-tong shape. Four rows of denticles
gomph falcigers. Both pairs of carriers and the four present.
basal plates of the jaws fused into a single structure.
Parophryotrocha Hartmann-Schroder 1971, Ophryo-
Dorvillea Parfrtt 1866, Staurocephalus rubrovittatus trocha isochaeta Eliason 1962; only species.
Grube 1855; 15 species. Dorvilleids with antennae and palps absent. Dorsal
Dorvilleids with both antennae and palps well de- cirri absent. Only simple, acicular setae present. Four
veloped and of the same length. Dorsal cirri with rows of denticles present, carriers and basal plates
acicula. Setae include capillaries and heterogomph fused into one piece.
falcigers. Carriers and four rows of denticles always
present; carriers may be variously fused to the basal Protodorvillea Pettibone 1961, Staurocephalus kefer-
plates. steini McIntosh 1869; 5 species.
Exallopus Jumars 1974, E. cropion Jumars 1974; only

species.
Dorvilleids with both antennae and palps well de-
veloped and of the same length. Dorsal cirri absent.
Setae include capillaries and composite heterogomph
falcigers and in the first setiger, large, curved com-
posite hooks. One pair of carriers and two rows of
denticles present; carriers fused to the basal plates.
Meiodorvillea Jumars 1974, Protodorvillea minuta
Hartman 1965; 3 species.
Dorvilleids with small clavate antennae, and small
palps, which may be absent. Dorsal cirri small or
absent, never with acicula. Setae include capillaries,
furcate setae and composite heterogomph falcigers. FIGURE 32. Family DORVILLEIDAE, Dorvillea articulara,
Carriers and two rows of denticles present. Newport, California, shallow subtidal, 50x.

Dorvilleids with well-developed palps; antennae small Eteonopsis Esmark 1878, see Ophryotrocha
or absent. Dorsal cirri without acicula. Setae include Prionognathus Keferstein 1862, see Schistomeringos
capillaries, furcate setae and composite heterogomph Staurocephalus Grube 1855, see Dorvillea
setae. Carriers and four rows of denticles present; car- Stauroceps Verrill 1900, see Dorvillea
riers sometimes fused with basal plates. Stauronereis Verrill 1900, see Dorvillea and Schisto-
meringos
Schistomeringos Jumars 1974, Nereis rudolphii delle
Telonereis Verrill 1900, see Dorvillea
Chiaje 1828; 10 species.
Dorvilleids with well-developed antennae and palps
The following two families are considered here
of approximately the same length. Dorsal cirri with
free-standing, unrelated families of the order Eunicida.
acicula. Setae include capillaries, furcate setae and
They are both very small, in terms of numbers of species,
composite heterogomph falcigers. Four rows of den-
and parasitic on decapod crustaceans and fishes, re-
ticles present; carriers may be fused with basal plates.
spectively. Probably in response to this habit, they
have been modified so that the only characters they
Taxonomic Notes
have in common with other members of the order, is
The genera of this family have been confused to a the structure of the jaw-apparatus, which makes them
considerable extent, as noted by Pettibone (1961) and resemble members of the Eunicida more closely than
Jumars (1974). Thanks to these two surveys, the matter they resemble members of any other jawed family.
seems to have been adequately clarified.
FAMILY HISTRIOBDELLIDAE VAILLANT 1890
Invalid Genera
Eunicida with five antennae; one pair of lower jaws
Anisoceras Grube 1856, see Dorvillea and indeter- and a single lateral jaw. One pair of anterior and one
minable pair of posterior appendages always present; a varying
e
FIGuRE 33. (A), Family ICHTHYOTOMIDAE, (chthyotomus sanguinarius, combined from several sources, ventral view, about
30x; (B), Family HISTRIOBDELLIDAE, Histriobdella homari, from Homarus americanus, Woods Hole, Massachusetts, 95x;
(C), Family STERNASPIDAE, Sternaspis scutata, off Santa Catalina Island, 23 m, anterior end inverted, 5x; (D), the above
with the anterior end everted, lox.

number of lateral appendages also present. Setae ab- marine environments. They are very small, and that,
sent. Parasitic on reptant decapod crustaceans. combined with their habitat makes it likely that they
Histriobdellids are known as parasites in the branchial are considerably more common than the few scattered
chambers of crustaceans, both from fresh-water and records indicate.
Key to Genera
Ia. With a single pair of lateral appendages Histriobdella

l b. With at least two pairs of lateral appendages Stratiodrilus
The genera are defined as indicated in the key. Each Renier 1807. The total number of currently recognized
genus is known in just a few species. species is about ten.
The sternaspids are among the most easily recognized
Histriobdella van Beneden 1858, H. homari van
polychaetes with the usually dark yellow or reddish
Beneden 1858, only species.
chitinized shield. They are common in sandy and
Stratiodrilus Haswell 1900, S. tasmanicus Haswell muddy substrates in all depths, but are perhaps most
1900, 4 species. usually found in about 100-200 m depth. Sternaspids
are only rarely found in large numbers. They are bur-
Invalid Genus rowers in the sand and mud.
Histriodrilus Foettinger 1884, see Histriobdella
Invalid Genera
FAMILY ICHTHYOTOMIDAE EISIG 1906 Echinorhynchus SENsu Renier 1804, see Sternaspis
Schreiberius Otto 1821, see Sternaspis
Eunicida with one antenna, one pair of lateral jaws.
Thalassenut Ranzani 1817, see Sternaspis
Notopodia with acicula, but otherwise asetigerous,
neurosetae composite. Parasitic on fishes. ORDER OWENIIDA
(chthyotomus is known for a single species, I.
sanguinarius Eisig 1906 from the Gulf of Naples. Prostomium fused to the anterior segments; pros-
It is parasitic on the fins of eels. Fauvel (1958) gives tomium sometimes produced in lobes or as a folded
a good description and illustration of the form; the membrane; proboscis a muscular pad. Neuropodial
illustration given here has been redrawn from that hooks in dense fields.
illustration.
FAMILY OWENIIDAE RIOJA 1917
ORDER STERNASPIDA Body cylindrical with long anterior segments and
Posterior ventrum covered by a stiff, chitinized, short posterior ones; tubicolous. Notosetae capillary,
mineral-impregnated shield. Eversible pharynx axial, neurosetae very small bi- or tridentate hooks in dense
can be inverted with the first three setigers. fields.
The oweniids are characteristically rather small,
tubicolous animals, the tubes are usually short, and
FAMILY STERNASPIDAE CARUS 1863
they are often capable of moving around with the tube.
Short-bodied polychaetes with indistinct segmenta- They have turned out to be quite frequently reported
tion; prostomium without appendages. All setae simple, from moderate depths on the continental slopes, but
those in the first three setigers thick, falcate spines; do not appear to be common in abyssal depths. The
those associated with the shield, slender capillaries. shape of the rather characteristic small hooks has been
The family is known for one genus, Sternaspis Otto well demonstrated by Thomassin and Picard (1972)
1821, with type species, Echinorhynchus scutatus with help of scanning electron microscope.
Key to Genera
la. Prostomium anteriorly produced into a low collar or tentacular crown 2
lb. Prostomium rounded or bilobed 3
2a (Ia). Prostomium forming a low collar, ventrally deeply incised Galathowenia
2b (Ia). Prostomium forming a tentacular crown, ventrally entire Owenia
3a (I b). Prostomium deeply bilobed with paired palps Myriowenia

3b (lb). Prostomium rounded, palps absent 4

4a (3b). First two or three setigers with notosetae only Myriochele
4b (3b). Only first setiger with notosetae only Myrioglobula
Galathowenia Kirkegaard 1959, G. africana Kirke-

gaard 1959; only species.
Prostomium anteriorly produced into a low collar that
ventrally is deeply incised with one lobe overlapping
the other. Eyes present. First three setigers without
neurosetae.
Myriochele Malmgren 1867, M. heeri Malmgren 1867;

12 species.
Prostomium anteriorly rounded. First two or three
setigers with notosetae only.
Myrioglobula Hartman 1967, M. antarctica Hartman

1967; only species.
Prostomium anteriorly rounded. Only one anterior
setiger with notosetae only.
Myriowenia Hartman 1960, M. californiensis Hartman
1960; 2 species.
Prostomium deeply bilobed with a pair of grooved
palps attached anteriorly. First three setigers without
neurosetae.
Owenia delle Chiaje 1841, O. fusiformis delle Chiaje

1841; 11 species.
Prostomium produced anteriorly into a low tentacular
crown with flattened lobate projections. Three first
setigers without neurosetae.
Invalid Genera
Ammochares Grube 1847, see Owenia

Mitraria Muller 1851, larval forms
Ops Carrington 1865, see Owenia
Psammocollus Grube 1868, see Myriochele
ORDER FLABELLIGERIDA
Anterior pharynx with ventrolateral muscular pad or

unreinforced. Anterior end retractable within a sheath
formed by the first setigers.
FAMILY FLABELLIGERIDAE SAINT-JOSEPH 1894

Body cylindrical or fusiform, most forms with epi-
FIGURE 34. (A), Family OWENIIDAE, Owenia collaris, com- dermis covered by papillae. Pro- and peristomia re-
bined from several sources. 12.5x; (B), Family FLABELLI- tractable between the first three setigers. Prostomium
GERIDAE, Pherusa inflata, after Hartman, 1969, 5x; (C), a slender ridge with palps at the posterolateral sides.
Family POEOBIIDAE, Poeobius meseres, redrawn and sim- Peristomium with an expanded dorsal membrane carry-
plified after Robbins, 1965, about 5x; (D), Family FAUVELI-
OPSIDAE, Fauveliopsis brevis, after Hartman and Fauchald, ing branchiae. Notosetae cross-barred or smooth, and
1 971, 12x. slender; neurosetae either similar or more thickened

spines or composite with falcate, unidentate or bidentate individual papilla, which secretes the mucus, has its
appendages. separate cover of particles. Dissection of the retractable
The flabelligerids characteristically are heavily im- anterior end is necessary for safe identification of the
pregnated with sand or mud in a matrix of mucus; in several similar genera. The number and structure of
some forms this mucus forms a complete smoothen- the branchiae and the structure of the branchial mem-
casing for the animal (Flabelligera), but in most each brane are important identificatory characters.
Key to Genera
Ia. Body with a distinct incision just posterior to the setigers carrying the cage-forming setae
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ......................................... Therochaeta
1b. Body without distinct incisions 2
2a (lb). Neurosetae composite or pseudocomposite 3
2b (lb). Neurosetae entirely simple, but usually distinctly cross-barred 4
3a (2a). Body encased in a smooth continuous mucus sheath Flabelligera
3b (2a). Body with individual papillae covered with mucus and impregnated with debris Flabelliderma
4a (2b), Branchiae absent Bradabyssa
4b (2b). Branchiae present 5
5a (4b). Branchial membrane long, tonguelike, sometimes doubled 6
5b (4b). Branchial membrane short, rounded or triangular 7
6a (5a). Branchial membrane club-shaped, with branchial filaments attached distally on all sides .. Coppingeria
6b (5a). Branchial membrane flattened, with branchial filaments attached on one side only Piromis
7a (5b). All setae capillary 8
7b (5b). At least some neurosetae acicular or falcigerous 9
8a (7a). Body anteriorly inflated with tapering posterior end Diplocirrus
8b (7a). Body, short, flattened and nearly disc-shaped Ilyphagus
9a (7b). Cephalic cage poorly developed or absent 10
9b (7b). Cephalic cage well developed 11
l0a (9a). Neurosetae distinctly thicker than notosetae, four pairs of branchiae Trophoniella
l0b (9a). Neurosetae only slightly thicker than notosetae; numerous pairs of branchiae Brada
I la (9b). Body covered by a thick mucus sheath Buskiella
lib (9b). Body not covered by a mucus sheath, often sand-incrusted 12
12a (11 b). A long oral tube present Therochaetella
l2b (Ilb). Oral tube absent 13
13a (12b). Notosetae serrated and plumose Pantoithrix
13b (12b). Notosetae cross-barred capillaries Pherusa
Generic Definitions prominent. Oral tube present. Branchial membrane

triangular with numerous branchial filaments. Neuro-
Brada Stimpson 1854, B. granola Stimpson 1854;
21 species. setae acicular.
Body short and fusiform; cephalic cage absent. Coppingeria Haswell 1892, C. longisetosa Haswell
Numerous branchial filaments on a short branchial 1892; only species.
membrane. Neurosetae slightly thicker than notosetae, Body anteriorly inflated; cephalic cage present.
distally acicular. Branchial membrane long, cylindrical and slender,
Bradabyssa Hartman 1967, B. papillata Hartman 1967; with the branchial filaments attached on all sides dis-
only species. t ally. Neurosetae simple hooks.
Body short and fusiform; cephalic cage absent. Diplocirrus Haase 1915, Trophonia glauca Malmgren
Branchiae absent; branchial membrane short and folded. 1867; 7 species.
Neurosetae simple hooks. Body anteriorly inflated; cephalic cage present in
Buskiella McIntosh 1885, B. abyssorum McIntosh some forms. Four pairs of branchiae of two kinds on
1885; 2 species. a short branchial membrane. All setae capillaries.
Body anteriorly inflated; mucus sheath covering the Flabelliderma Hartman 1969, Flabelligera commensalis
whole body. Cephalic cage present; parapodia very Moore 1909; only species.

Body cylindrical; body papillae covered with thick Invalid Genera

mucus and encrusted with debris. Cephalic cage present.
Aristenia Savigny in Quatrefages 1865, indeterminable
Pseudocomposite hooks present in most neuropodia.
Balanochaeta Chamberlin 1919c, see Pherusa
Flabelligera Sars 1829, F. affinis Sars 1829; 18 Chloraema Dujardin 1839a, see Flabelligera
species. Flemingia Johnston 1846, see Pherusa
Body cylindrical; covered completely by smooth Lophocephalus Costa 1841, see Pherusa
mucus sheath. Cephalic cage present; pseudocomposite Pycnoderma Grube 1877, see Piromis
or composite hooks in the neuropodia. Saphobranchia Chamberlin 1919c, see Diplocirrus
Semiodera Chamberlin 1919c, see Piromis
Ilyphagus Chamberlin 1919c, I. bythincola Chamberlin Siphonostoma Rathke 1843, see Flabelligera
1 919c; 11 species. Siphostoma Otto 1821, see Flabelligera
Body stout, flattened and nearly disc-shaped, covered Stylarioides delle Chiaje 1841, see Pherusa
with large papillae and mud- or sand-incrusted in most Tecturella Stimpson 1854, see Flabelligera
forms. Cephalic cage present in some forms. Four pairs Trophonia Cuvier 1830, see Pherusa
of branchiae on a short branchial membrane. All setae Zorus Webster and Benedict 1887, indeterminable
capillaries.
FAMILY POEOBIIDAE HEATH 1930
Pantoithrix Chamberlin 1919c, Pherusa chilensis
Schmarda 1861; only species. Body saclike without external segmentation or setae.
Body anteriorly inflated; cephalic cage present. Noto- Anterior end fused to the rest of the body; containing
setae serrated and plumose capillaries; neurosetae bi- a circlet of eversible tentacles. Two distinct septa only
dentate hooks. Six pairs of branchiae. polychaete characters. Pelagic.
The poeobiids are considered related to the flabel-
Pherusa Oken 1807, Amphitrite plumosa O.F. MUller ligerids because the anterior end is retractable, and
1 776; 43 species. because they have chlorocruorin as one of their pig-
Body anteriorly inflated; cephalic cage present. Either ments (Robbins 1965). The family contains a single
four or many pairs of branchiae on a short branchial genus and species, Poeobius meseres Heath 1930.
membrane. Most neurosetae uni- or bidentate hooks.
Piromis Kinberg 1867b, P. arenosus Kinberg 1867b; Incertae Sedis

1 0 species. Enigma Betrem 1924, E. terwillei Betrem 1924; only
Body anteriorly inflated; cephalic cage present. species.
Branchial membrane prolonged, flattened and tongue- Parapodia and setae absent, retractile branchiae and
shaped, either single or double, with numerous branchial palps present; one septum observed, pelagic.
filaments. Neurosetae mostly uni- or bidentate hooks. This form resembles the poeobiids in several re-
spects, it has been reported only once, but unfortunately
Therochaeta Chamberlin 1919c, Srylarioides collarifer
the original material has been lost (Hartman 1967).
Ehlers 1887; 6 species.
Anterior end inflated; cephalic cage present. A dis-
ORDER FAUVELIOPSIDA
tinct incision present behind the last setiger that carries
the cage setae. First post-incisional setiger with series Pro- and peristomium without appendages. Pharynx
of enlarged, usually anteriorly directed papillae. Com- with a ventral muscular pad. All setigers biramous
posite hooks present on some anterior setigers in most with simple limbate setae and a small rounded papilla
species; otherwise simple neuropodial hooks. between the rami.
These small, deep-water polychaetes are incom-
Therochaetella Hartman 1967, T. chilensis Hartman pletely known. They were grouped formerly with the
1967; only species. flabelligerids from which they differ in that they lack
Anterior end inflated; cephalic cage present. Long the papillar investments, the retractable anterior end
narrow oral tube present. Numerous pairs of branchiae and the characteristic setae. The shape of the pros-
on a short branchial membrane. Neurosetae distally tomium also is markedly different. Further information
bifid and falcate. may demonstrate that they are related to other groups
Trophoniella Caullery 1944, T. avicularia Caullery of polychaetes; a separate order appears justified for
1944; 3 species. the time being.
Anterior end inflated; cephalic cage very poorly de-
veloped or absent. Four pairs of branchiae on a short FAMILY FAUVELIOPSIDAE HARTMAN 1971
branchial membrane. Neurosetae bi- or unidentate Small, smooth-bodied polychaetes without anterior
hooks. appendages. Proboscis a ventral muscular pad. All

setigers biramous with simple limbate setae and a small lobed processes. Pharynx muscular and eversible,
rounded papilla between the rami. Parapodial lobes scoop-shaped and open dorsally. Parapodia biramous
reduced. with setae of two kinds; slender smooth acicular rods
The family was erected by Hartman (1971) for and thicker and cross-barred acicular setae. Body with
Fauveliopsis McIntosh, with type F. challengeriae papillae, especially on the parapodia. Encased in a
McIntosh 1922, as well as for Flabelligella Hartman thick mucus sheath, pelagic.
1965, Flota Hartman 1967 and Bruunilla Hartman
1971. Flabelligella was shown to belong to the Acro- ORDER TEREBELLIDA
cirridae by Orensanz (1974b) and are cited under that
Prostomium without appendages; peristomium with
family above. Flora and Bruunilla differ sharply from
a series of feeding appendages; pharynx with a ventral
Fauveliopsis and are characterized here as free-standing
muscular pad. At least one pair of branchiae present
genera without obvious familial affiliations.
(rarely absent).
The whole family is then reduced to the type-genus,
Fauveliopsis, with eight described species.
FAMILY SABELLARIIDAE JOHNSTON 1 865
Incertae Sedis Tubicolous polychaetes with the body in three re-

gions; posterior region an asetigerous anal tube. Pros-
Bruunilla Hartman 1971, B. natalensis Hartman 1971;
tomium a narrow ridge fused laterally to the first setiger.
only species.
Setae of first setiger forming an operculum with setae
Prostomium small and triangular attached ventral
in one to three rows. Thorax with two rudimentary
to the large peristomium. Peristomium with five ten-
segments and three or four parathoracic setigers.
tacular cirri. Paired short palps lateral to the mouth as
Median region with notopodial pectinate uncini and
flat pads. Prominent lateral parapodia present; all bi-
ventral capillary setae.
ramous, both rami distally strongly pointed, with
The sabellariids are important as reef-builders in
embedded acicula, setae absent. Dorsal and ventral
certain areas (Schafer 1972). All of them are tube-
cirri present on most segments.
builders and tend to build on firm substrates; conse-
Flota Hartman 1967, F. flabelligera Hartman 1967; quently, most of them are present in shallow water
only species. only, but members of two genera (Phalacrostemma
Body with less than ten segments; short and flattened. and Monorchos) appear to be most common in slope
Prostomium a simple conical lobe with a pair of tri- depths.
Key to Genera
Ia. With a single row of paleae in the operculum 2

l b. With at least two rows of paleae in the operculum 3
2a (1 a). The prolonged opercular peduncles free from one another Phalacrostemma
2b (la). Opercular peduncles short and fused Monorchos
3a (lb). Two rows of paleae 4
3b (I b). Three rows of paleae 6
4a (3a). Hooks absent Gunnarea
4b (3a). A pair of large hooks dorsal and proximal to the opercular paleae 5
5a (4b). Three parathoracic setigers Idanthyrsus
5b (4b). Four parathoracic setigers Lygdamis
6a (3b). Middle opercular paleae cover the inner ones; operculum flattened cone Phragmatopoma
6b (3b). Middle opercular paleae not concealing the inner ones; operculum open and bristly Sabellaria
Generic Definitions Two rows of paleae; opercular peduncles fused;

hooks present dorsal and proximal to the paleae; ac-
Gunnarea Johansson 1927, Hermella capensis Schmarda cessory setae absent. Three parathoracic setigers.
1861; only species.
Lygdamis Kinberg 1867b, L. indicus Kinberg 1867b;
Two rows of paleae; opercular peduncles fused;
12 species.
hooks and accessory setae absent on operculum. Three Two rows of paleae; opercular peduncles long and
parathoracic setigers.
separated; hooks present dorsal and proximal to the
Idanthyrsus Kinberg 1867b, I. armatus Kinberg 1867b; paleae; accessory setae absent. Four parathoracic
5 species. setigers.

LJ
A B
FIGuRE 35. (A), Family PECTINARIIDAE, Amphictene capensis, after Day, 1967, 3x; (B), pectinariid-tube, 2x; (C), Family
SABELLARIIDAE, tube pattern of Phragmatopoma sp. natural size; (D), Phragmatopoma californica, after Hartman, 1969, 5x.
Monorchos Treadwell 1926, M. philippinensis Tread- paleae; accessory setae absent. Four parathoracic
well 1926; 2 species. setigers.
A single row of paleae; opercular peduncles fused;
Phragmatopoma Morch 1863, P. caudata Morch 1863;
hooks present dorsal and proximal to the paleae and
8 species.
two rows of accessory setae between the paleal rows.
Three rows of paleae; opercular peduncles short and
Three parathoracic setigers.
fused; hooks and accessory setae absent. Three para-
Phalacrostemma Marenzeller 1895, P. cidariophilam thoracic setigers. Middle opercular paleae covers the
Marenzeller 1895; 5 species. inner ones; operculum conical.
A single row of paleae; opercular peduncles long Sabellaria Savigny 1818, Sabella alveolata Linnaeus
and separated; hooks present dorsal and proximal to the 1767; 28 species.

Three rows of paleae; opercular peduncles short and Eupallasia Augener 1927a, see Lygdamis
fused; hooks absent, accessory setae sometimes present. Hermella Savigny 1818, see Sabellaria
Three parathoracic setigers. Middle opercular paleae Pallasia Quatrefages 1848, see Idanthyrsus
pointed distad, operculum open and generally rather Pallasina Annenkova 1925, see Idanthyrsus
bristly in appearance. Tetreres Caullery 1913, see Lygdamis
Taxonomic Notes 1865

FAMILY PECTINARIIDAE QUATREFAGES
The "accessory setae" referred to in the above defi-
Body separated into three regions, including thorax,
nitions, include setae associated with the operculum;
abdomen and a posterior scaphe. Prostomium reduced.
they usually are acicular in appearance and may be
First setiger with expanded, strong setae (paleae) form-
present between the rows of paleae (Monorchos) or
ing a comb used in digging. Other setae include short
proximal to the paleae proper (Sabellaria). The family
capillaries, pectiniform uncini and scaphal spines.
is under current revision by Dr. David Kirtley and Tubes short, more or less tusk-shaped, built of rela-
the numbers of species assigned to each genus may be
tively large particles.
changed drastically. The generic sub-division appears
The pectinariids are among the most characteristic
reasonably stable.
polychaetes with their strongly golden, coppery or
brassy paleal setae and the gently curved, tapering,
Invalid Genera
tusk-shaped, but usually very fragile tubes. The family
Centrocorone Grube 1850, see Sabellaria often is recognized for only two genera, Petta and Pec-
Chrysodon Oken in Quatrefages 1865, see Sabellaria tinaria, in which case the other listed genera are con-
Cryptopomatus Gravier 1908, see Idanthyrsus sidered subgenera of Pectinaria.
Key to Genera
la. Cephalic veil marginally smooth, scaphe not distinctly separated from abdomen Petta
lb. Cephalic veil marginally cirrate; scaphe distinctly set off from abdomen 2
2a (I b). Twelve uncinigers, cephalic veil at least partly fused to the operculum Lagis
2b (lb). Thirteen uncinigers, cephalic veil completely free from the operculum 3
3a (2b). Opercular rim cirrate Amphictene
3b (2b). Opercular rim smooth 4
4a (3b). Uncini with major teeth in a single row Cistenides
4b (3b). Uncini with major teeth in two rows Pectinaria
Generic Definitions distinctly set off from abdomen. Uncini with major
teeth in two or more rows; 12 uncinigers present.
Amphictene Savigny 1818, Amphitrite auricoma O.F. Pectinaria Savigny 1818, Nereis cylindraria belgica
Miiller 1776; 8 species. Pallas 1766; 18 species.
Cephalic veil free from operculum, marginally Cephalic veil free from operculum, marginally cir-
citrate; opercular rim marginally citrate. Scaphe rate. Opercular rim marginally smooth. Scaphe dis-
distinctly set off from abdomen. Uncini with major tinctly set off from the abdomen. Uncini with major
teeth in double rows; 13 uncinigers. teeth in double rows; 13 uncinigers present.
Cistenides Malmgren 1866, Sabella granulata Lin- Petta Malmgren 1866, P. pusilla Malmgren 1866;
naeus 1767; 8 species. 4 species.
Cephalic veil free from operculum, marginally Cephalic veil free from operculum; marginally
cirrate; opercular rim marginally smooth. Scaphe smooth. Opercular rim marginally cirrate. Scaphe
distinctly set off from abdomen. Uncini with major not distinctly set off from abdomen. Uncini with major
teeth in a single row, 13 uncinigers present. teeth in a single row; 14 uncinigers.
Lagis Malmgren 1866, L. koreni Malmgren 1866; Invalid Genera
8 species.
Cephalic veil at least partly fused to operculum, mar- Ariapithes Kinberg 1867b, indeterminable
ginally curate; opercular rim marginally smooth. Scaphe Cistena Leach 1816, see Pectinaria

Labiaria Sveshnikov 1959, larval forms ally, the ampharetids usually have a few pairs of simple
Scalis Grube 1846, indeterminable branchiae, never the masses of arborescent branchiae or
the numerous sessile filaments present in the terebellids.
The uncini often are flattened plates in the ampharetids
FAMILY AMPHARETIDAE MALMGREN 1867 and nearly always distinctly crested in the terebellids.
The ampharetids have turned out to be very common
Body with two regions; anterior region with biramous in deep water and a whole mass of previously unrec-
parapodia; posterior region with well-developed neu- ognized genera have been reported from deep water
ropodia, notopodia reduced or absent. Prostomium over the last ten years. The major classification was
simple or complex with lateral folds and glandular reviewed by Day (1964) who reduced greatly the num-
ridges. Two to four pairs of smooth or pinnate bran- ber of monotypic genera. The present review retains
chiae present. Notopodial capillary setae present in most of the genera Day fused; it is not clear that the
thorax, neuropodial uncini present both in thorax and characters Day used to identify his genera are any more
abdomen. Thoracic uncini with major teeth in one or a precise than those he disregarded. Since a large num-
few rows, rarely crested; abdominal uncini similar, but ber of additional taxa are now being described, it ap-
more frequently crested. Nuchal hooks and anterior pears best to await further fusions of genera, until the
acicular setae present in some forms. current deep-water material has been worked up in de-
The ampharetids resemble the terebellids; the main tail. The genus Oeorpata is not clearly separable from
feature used to separate the two families is behavioral in Isolda according to Day (1964) and is incompletely
that the former will withdraw the buccal tentacles com- known. It has been included in the definitions, but not
pletely within their mouth; the latter do not. Addition- in the key.
Key to Genera
Ia. First several neuropodia with fine acicular setae; other neuropodia with uncini MELINNINAE
2
l b. First several neuropodia without acicular setae; all neurosetae uncini AMPHARETINAE
10
2a (la). Nuchal hooks present 3
2b (Ia). Nuchal hooks absent 6
3a (2a). Branchiae smooth 4
3b (2a). Branchiae in part papillose or pennate 5
4a (3a). Two pairs of nuchal hooks; 12 thoracic uncinigers Moyanus
4b (3a). A single pair of nuchal hooks, 14 thoracic uncinigers Melinna
5a (3b). Four pairs of branchiae, two smooth and two papillose; capillary notosetae on setigers
3-4 Isolda
5b (3b). Three pairs of branchiae, one smooth, two papillose; notosetae absent on setigers 3-4
Iran
6a (2b). Ten thoracic uncinigers present 7
6b (2b). At least 12 thoracic uncinigers present 8
7a (6a). Three pairs of branchiae Melinnopsides
7b (6a). Four pairs of branchiae Melinnopsis
8a (6b). Sixteen thoracic uncinigers present; all buccal tentacles similar Melinantipoda
8b (6b). Maximally 14 thoracic uncinigers present; one or two buccal tentacles very large 9
9a (8b). Segment 6 with a distinct dorsal crest between the notopodia Melinnexis
9b (8b). A dorsal crest absent Amelinna
l0a (Ib). At least three pairs of branchiae 11
l0b (lb). Two pairs of branchiae 12
I la (l0a). Three pairs of branchiae present 15
1lb (l0a). Four pairs of branchiae 35
12a (l0b). Fourteen thoracic uncinigers; paleae present Ecamphicteis
12b (l0b). Maximally 12 thoracic uncinigers; paleae absent 13
13a (12b). Nine uncinigers, first uncini in a short row similar to all others Egamella
13b (12b). Twelve uncinigers, first uncini in a long row 14
14a (13b). Prostomium anteriorly pointed Auchenoplax

FIGuRE 36. (A), Family BOGUEIDAE, Boguella ornata, redrawn after Hartman and Fauchald, 1971, about 23x; (B), Family
AMPHARETIDAE, Amphicteis scaphobranchiata, modified after Fauchald, 1972, 12.5x; (C), Family TRICHOBRANCHIDAE,
Terebellides stroemi, combined from several sources, about 5x; (D), Family TEREBELLIDAE, Neoamphitrite, near johnstoni,
modified from live sketch, 3x.

14b (13b). Prostomium anteriorly truncate Melinnoides

l5a (11 a). All branchial pairs arranged in a distinctly segmental fashion 16
15b (I la). Only last pair of branchiae clearly associated with a specific segment 18
16a (15a). Eleven thoracic uncinigers; tentacular membrane large Amphora=
16b (15a). At least 14 thoracic uncinigers; tentacular membrane small 17
17a (16b). First setae on segment 2; 14 thoracic uncinigers Neopaiwa
17b (16b). First setae in segment 3; 15 thoracic uncinigers Weddellia
18a (15b). Paleae present 19
18b (15b). Paleae absent 25
19a (18a). Nine thoracic uncinigers; last thoracic notopodia elevated Mugga
l9b (18a). Ten or more thoracic uncinigers 20
20a (19b). Flange present between last thoracal notopodia; ten thoracic uncinigers Melinnata
20b (19b). Flange or crest, if present, anterior in position; at least I I thoracic uncinigers 21
2Ia (20b). Few stout buccal tentacles present; I1 thoracic uncinigers Amythasides
21b (20b). Buccal tentacles, if present, numerous, 12 thoracic uncinigers 22
22a (21b). Dorsal crest present on one anterior setiger 23
22b (21b). Dorsal crest absent Eclysippe
23a (22a). Oral membrane broadly folded, 15 thoracic setigers Eusamythella
23b (22a). Oral membrane short; 14 thoracic setigers 24
24a (23b). Dorsal crest on segment 4 Neosamytha
24b (23b). Dorsal crest on either segment 6 or 7 Melinnampharete
25a (18b). Ten thoracic uncinigers Muggoides
25b (18b). At least 11 thoracic uncinigers 26
26a (25b). Eleven thoracic uncinigers Glyphanostomum
27a (26b). Twelve thoracic uncinigers 28
28a (27a). At least one pair of branchiae basally flanged Samythella
28b (27a). All branchiae cylindrical 29
29a (28b). Buccal tentacles papillose Neosabellides
29b (28b). Buccal tentacles smooth 30
30a (29b). Oral membrane greatly prolonged Pabits
30b (29b). Oral membrane short Eusamytha
31a (27b). Thirteen thoracic uncinigers 32
31b (27b). Fourteen thoracic uncinigers 33
32a (31a). Notopodia of setiger 13 elevated with hirsute notosetae Sosanella
32b (31a). Notopodia of setiger 13 at the same level as all others; hirsute setae absent Alkmaria
33a (31b). Oral membrane smooth and folded Amythas
33b (31b). Oral membrane tentaculate 34
34a (33b). Glandular ridges on prostomium Samythopsis
34b (33b). Glandular ridges absent Samytha
35a (I lb). Paleae present (i.e. setae present on segment 3, as the first notosetae in the body) 36
35b (11b). Paleae absent (i.e. setae absent on segment 3 or setae already present from segment 2) 52
36a (35a). Flanged branchiae present on four successive segments Phyllampharate
36b (35a). Branchiae flanged, lamellate, pennate or cylindrical; only one pair clearly associated with a
segment 37
37a (36b). Eleven thoracic uncinigers 38
38a (37a). Branchiae pennate, buccal tentacles papillose Pterampharete
38b (37a). Both branchiae and buccal tentacles smooth Sabellides
39a (37b). Twelve thoracic uncinigers present 40
39b (37b). At least 13 thoracic uncinigers present 44
40a (39a). Buccal tentacles papillose 41
40b (39a). Buccal tentacles smooth 42
41a (40a). Setiger 11 with notopodia elevated and notosetae distally hirsute Anobothrella
41b (40a). Setiger I I with notopodia at the same level as all other setigers; notosetae limbate or capillary
. . . . . . . . . . . . . . . . . . . . . .....................................................Ampharete

42a (40b). Setiger 14 with elevated notopodia and hirsute notosetae Sosane
42b (40b). Setiger 10 or 11 with elevated notopodia and tapering or hirsute notosetae 43
43a (42b). Notopodial rudiments present in abdomen Anobothrus
43b (42b). Notopodial rudiments absent from abdomen Sosanides
44a (39b). Thirteen thoracic uncinigers : 45
45a (44a). One pair of branchiae pennate, others smooth Pterolysippe
45b (44a). All branchiae smooth 46
46a (45b). Glandular ridge on prostomium Hypania
46b (45b). Glandular ridge absent Lysippe
47a (44b). Two of the four pairs of branchiae lamellate 48
47b (44b). All four pairs of branchiae cylindrical 49
48a (47a). Paleae large, abdominal notopodial rudiments absent Phyllamphicteis
48b (47a). Paleae small, abdominal notopodial rudiments present Lysippides
49a (47b). Glandular ridges on prostomium 50
49b (47b). Glandular ridges absent 51
50a (49a). Abdominal notopodial rudiments present Amphicteis
50b (49a). Abdominal notopodial rudiments absent Parhypania
51a (49b). Abdominal notopodial rudiments present Paiwa
51b (49b). Abdominal notopodial rudiments absent Hypaniola
52a (35b). Branchiae arranged in oblique series directly associated with distinct segments Mexamage
52b (35b). Only the last pair of branchiae clearly associated with a segment 53
53a (52b). At least three pairs of branchiae lamellate 54
53b (52b). All branchiae cylindrical 55
54a (53a). All four pairs of branchiae lamellate; anus surrounded by a circle of papillae Phyllocomus
54b (53a). Three pairs of lamellate and one pair of cylindrical branchiae; anus with two pairs of anal cirri ....
Schistocomus
55a (53b). Eleven thoracic uncinigers 56
56a (55a). First two notopodia (segments 4 and 5) asetigerous Paramage
56b (55a). All anterior notopodia with setae 57
57a (56b). One pair of anal cirri Grubianella Y
57b (56b). Two pairs of anal cirri Amage
58a (55b). Twelve thoracic uncinigers 59
59a (58a). Buccal tentacles papillose; all notosetae capillary Asabellides
59b (58a). Buccal tentacles smooth; last thoracic notosetae modified Sosanopsis
60a (58b). Buccal tentacles papillose; notopodial cirri present Paramphicteis
60b (58b). Buccal tentacles smooth; notopodial cirri absent Amphisamytha
Generic Definitions MELINNINAE. Four pairs of smooth branchiae;

notosetae absent in segments 3 and 4. Nuchal hooks
Alkmaria Horst 1919b, A. rominji Horst 1919b; only absent; one or two large oral tentacles present in
species.
addition to numerous smaller ones. Twelve or 13
AMPHARETINAE. Three pairs of smooth branchiae;
thoracic uncinigers.
no paleae. Glandular ridges absent; 13 thoracic un-
cinigers; no notopodial rudiments in abdomen. Ampharana Hartman 1967, A. antarctica Hartman
Amage Malmgren 1866, A. auricula Malmgren 1866; 1967; only species.
14 species. AMPHARETINAE. Three pairs of smooth branchiae
AMPHARETINAE. Four pairs of smooth branchiae; on three successive segments. Oral membrane large
no paleae. Glandular ridges present; 11 thoracic un- and folded with buccal tentacles in two lateral groups.
cinigers; abdominal notopodial rudiments present. Eleven thoracic uncinigers. Paleae absent, but small
Two pairs of anal cirri. capillary setae present in segment 3.
Amelinna Hartman 1969, A. abyssalis Hartman 1969; Ampharete Malmgren 1866, Amphicteis ac ifrons
2 species. Grube 1860; 27 species.

AMPHARETINAE. Four pairs of smooth branchiae; sent in segments 3 and 4. Twelve thoracic uncinigers;
paleae present. Buccal tentacles papillose. Glandular first row of uncini very long and ventrally located.
ridges absent. Twelve thoracic uncinigers; abdominal
Ecamphicteis Fauchald 1972, E. elongata Fauchald
notopodial rudiments absent. Notosetae absent in
1972; only species.
segment 4.
AMPHARETINAE. Two pairs of smooth branchiae
Amphicteis Grube 1850, Amphitrite gunneri Sars 1835; on first two segments. Paleae present; 14 uncinigers.
30 species. Glandular ridges and abdominal notopodial rudiments
AMPHARETINAE. Four pairs of branchiae, usually absent.
cylindrical, rarely foliose; paleae present. Glandular
Eclysippe Eliason 1955, Lysippe vanelli Fauvel 1936;
ridges present. Fourteen thoracic uncinigers; abdominal
only species.
notopodial rudiments present.
AMPHARETINAE. Three pairs of smooth branchiae
Amphisamytha Hessle 1917, A, japonica Hessle 1917; (Day 1964: four pairs of branchiae in type species);
2 species. paleae present; 12 thoracic uncinigers. Glandular ridges
AMPHARETINAE. Four pairs of smooth branchiae; absent; abdominal notopodial rudiments absent.
paleae absent. Glandular ridges absent. Fourteen tho-
Egamella Fauchald 1972, E. quadribranchiata Fau-
racic uncinigers. Abdominal notopodial rudiments
chald 1972; only species.
present.
AMPHARETINAE. Two pairs of branchiae on two
Amythas Benham 1912, A. membranifera Benham successive segments; branchial membrane high and
1921; only species. laterally free from branchial bases. Segment 3 with
AMPHARETINAE. Three pairs of smooth or grooved small capillary setae; nine uncinigers present. Clavate
branchiae; oral membrane folded and smooth. Palette notopodial abdominal rudiments present; glandular
absent; 14 thoracic uncinigers. Abdominal notopodial ridges absent.
rudiments absent.
Eusamytha McIntosh 1885, E. pacifica McIntosh 1885;
Amythasides Eliason 1955, A. macroglossus Eliason only species.
1955; only species. AMPHARETINAE. Three pairs of smooth branchiae;
AMPHARETINAE. Three pairs of branchiae; paleae paleae absent. Glandular ridges and abdominal noto-
present. Buccal tentacles few and large. Eleven thoracic podial rudiments absent. Twelve thoracic uncinigers.
uncinigers. Glandular ridges absent. Abdominal noto-
podial rudiments absent. Eusamythella Hartman 1971, Eusamytha sexdentata
Hartman 1967; only species.
Anobothrella Hartman 1967, Anobothrus antarctica AMPHARETINAE. Three pairs of smooth branchiae;
Monro 1939a; only species. paleae present. Oral membrane broad and folded; dorsal
AMPHARETINAE. Four pairs of papillose branchiae. crest across segment 5. Twelve thoracic uncinigers.
Buccal tentacles papillose; paleae present. Twelve Glandular ridges absent.
thoracic uncinigers. Setiger 11 with notopodia elevated
and hirsute notosetae. Glyphanostomum Levihsen 1884, Samytha pallescens
Theel 1879; 2 species.
Anobothrus Levinsen 1884, Ampharete gracilis Malm- AMPHARETINAE. Three pairs of smooth branchiae;
gren 1866; 8 species. paleae absent. Glandular ridges absent. Eleven thoracic
AMPHARETINAE. Four pairs of smooth branchiae, uncinigers.
paleae present. Twelve uncinigers. Glandular ridges
absent; abdominal notopodial rudiments present. Setiger Grubianella McIntosh 1885, G. antarctica McIntosh
10 or 11 with notopodia elevated and modified notosetae. 1885; only species.
AMPHARETINAE. Four pairs of smooth branchiae;
Asabellides Annenkova 1929, Sabellides sibirica Wien paleae absent. Glandular ridges present; abdominal
1883; 3 species. notopodial rudiments present; one pair of anal cirri.
AMPHARETINAE. Four pairs of smooth branchiae; Eleven thoracic uncinigers.
paleae absent. Twelve thoracic uncinigers; notosetae
absent in segment 4. Buccal tentacles papillose; Hypania Ostroumouw 1897, Amphicteis i alida
glandular ridges absent. Abdominal notopodial rudi- Grube 1860; 2 species.
ments present. AMPHARETINAE. Four pairs of smooth branchiae,
paleae present. Glandular ridges and abdominal noto-
Auchenoplax Ehlers 1887, A. crinita Ehlers 1887;
podial rudiments present. Thirteen thoracic uncinigers.
only species.
AMPHARETINAE. Two pairs of smooth branchiae. Hypaniola Annenkova 1927, Amphicteis (?Aryandes)
Prostomium anteriorly sharply pointed. Notosetae ab- kowalewskii Grimm 1877; only species.

AMPHARETINAE. Four pairs of smooth branchiae; Melinnexis Annenkova 1931, M. arctica Annenkova
small paleae present. Glandular ridges absent; 14 1931; 8 species.
uncinigers. MELINNINAE. Four pairs of smooth branchiae.
Nuchal hooks absent; first notosetae in segment 5.
Irana Wesenberg-Lund 1949, 1. heterobranchiata
Dorsal crest on segment 6. Thirteen or 14 thoracic
Wesenberg-Lund 1949; only species.
uncinigers. One very large and numerous small buccal
MELINNINAE. Three pairs of branchiae, one
tentacles present.
smooth, other two pennate. Nuchal hooks on segment
4; dorsal crest on segment 6. Capillary notosetae Melinnoides Benham 1927, M. nelsoni Benham 1927;
first present in segment 7; 12 thoracic uncinigers. only species.
AMPHARETINAE. Two pairs of smooth branchiae;
Isolda Muller 1858,1. pulchella Muller 1858; 4 species.
paleae absent; first notosetae in segment 5. Prostomium
MELINNINAE. Four pairs of branchiae; two smooth a small quadrangular lobe. Twelve thoracic uncinigers;
and two pennate. Nuchal hooks present; dorsal crest first row of uncini long and ventrally displaced.
on segment 6. Capillary notosetae present in segments
5 and 6. Twelve or 13 thoracic uncinigers. Melinnopsides Day 1964, Melinnopsis capensis Day
1955; only species.
Lysippe Malmgren 1866, L. labiata Malmgren 1866; MELINNINAE. Three pairs of smooth branchiae;
5 species. nuchal hooks and dorsal crest absent. Ten thoracic
AMPHARETINAE. Four pairs of smooth branchiae; uncinigers. First notosetae in segment 5; segment 6
small paleae present. Glandular ridges absent. Ab- without neurosetae.
dominal notopodial rudiments present. Thirteen tho-
racic uncinigers. Melinnopsis McIntosh 1885, M. atlantica McIntosh
1885; only species.
Lysippides Hessle 1917, Amphicteis fragilis Wollebaek MELINNINAE. Four pairs of smooth branchiae;
1912; only species. nuchal hooks absent; first notosetae on segment 5.
AMPHARETINAE. Four pairs of branchiae; two Dorsal crest absent. Ten thoracal uncinigers. Buccal
cylindrical, two flanged; small paleae present. Glan- tentacles all similar.
dular ridges absent. Abdominal notopodial rudiments
present. Fourteen thoracic uncinigers. Mexamage Fauchald 1972, M. corrugata Fauchald
1972; only species.
Melinantipoda Hartman 1967, M. antarctica Hartman AMPHARETINAE. Four pairs of branchiae on four
1 967; only species. successive segments. Paleae absent; notopodia on seg-
MELINNINAE. Four pairs of smooth branchiae; ments 2 and 3, but not setae. Ten thoracic uncinigers;
nuchal hooks absent. First notosetae in segment 5. no fused anterior segments.
Dorsal crest on segment 6; 16 thoracic uncinigers. Moyanus Chamberlin 1919c, M. explorans Chamberlin
All buccal tentacles similar.
1919c; only species.
Melinna Malmgren 1866, Sabellides cristata Sars MELINNINAE. Four pairs of smooth branchiae.
1851; 26 species. Nuchal hooks on both segments 4 and 5. Dorsal crest
MELINNINAE. Four pairs of branchiae, nearly on segment 6. Twelve thoracic uncinigers. Prostomium
always smooth. Nuchal hooks present; dorsal crest prolonged with oral lobe suspended below it.
on segment 6 present. Fourteen thoracic uncinigers. Mugga Eliason 1955, M. wahrbergi Eliason 1955;
All buccal tentacles similar. only species.
Melinnampharete Annenkova 1937, M. eon Annenkova AMPHARETINAE. Three pairs of smooth branchiae;
1 937; 2 species. paleae present. Glandular ridges absent; abdominal noto-
AMPHARETINAE. Three pairs of smooth branchiae; podial rudiments absent. Nine thoracic uncinigers. Last
paleae present. Dorsal ridge either on segment 6 or thoracic notopodia elevated with modified notosetae.
7. Glandular ridges absent; abdominal notopodial Muggoides Hartman 1965, M. cinctus Hartman 1965;
rudiments absent. Twelve thoracic uncinigers. only species.
Melinnata Hartman 1965, M. americana Hartman AMPHARETINAE. Three pairs of smooth branchiae;
1965; only species. palette absent. Glandular ridges and abdominal noto-
AMPHARETINAF. Three pairs of branchiae; palette podial rudiments absent. Ten thoracic uncinigers; last
present. Glandular ridges absent; ridge across dorsum thoracic notopodia elevated with modified notosetae.
on segment 4. Ten thoracic uncinigers; flange across Neopaiwa Hartman and Fauchald 1971, N. cirrata
dorsum between last thoracic notopodia. Hartman and Fauchald 1971; only species.

AMPHARETINAE. Three pairs of branchiae on AMPHARETINAE. Four pairs of smooth branchiae;

three successive segments starting on segment 2. Cap- paleae present. Glandular ridges present; abdominal
illary notosetae present from segment 2; paleal devel- notopodial rudiments very small. Fourteen thoracic
opment absent. Fourteen thoracic uncinigers present. uncinigers.
Glandular ridges absent; abdominal notopodial rudi- Phyllampharete Hartman and Fauchald 1971, P.
ments well developed. longicirrata Hartman and Fauchald 1971; only
Neosabellides Hessle 1917, N. elongata Hessle 1917; species.
2 species. AMPHARETINAE. Four pairs of flanged branchiae
AMPHARETINAE. Three pairs of smooth branchiae; on four successive segments starting on segment 2.
paleae absent; first notosetae present on segment 5. Paleae absent; notosetae present from segment 2.
Glandular ridges absent; abdominal notopodial rudi- Glandular ridges absent; abdominal notopodial rudi-
ments present. Twelve thoracic uncinigers. Buccal ments present. Eleven thoracic uncinigers. Prostomium
tentacles papillose. anteriorly incised.
Phyllamphicteis Augener 1918, P. collaribranchis

Neosamytha Hartman 1967, N. gracilis Hartman 1967;
only species. Augener 1918; 2 species.
AMPHARETINAE. Three pairs of smooth branchiae; AMPHARETINAE. Four pairs of branchiae, two
paleae present. Glandular ridges and abdominal noto- smooth and two lamellate. Paleae present. Glandular
podial rudiments absent. Twelve thoracic uni inigers. ridges and abdominal notopodial rudiments absent.
Dorsal ridge across segment 4. Fourteen thoracic uncinigers.
Phyllocomus Grube 1877, P. crocea Grube 1877; only
Oeorpata Kinberg 1867b, O. armata Kinberg 1867;
species.
only species.
MELINNINAE. Incompletely known, not clearly AMPHARETINAE. Four pairs of flanged branchiae;
paleae absent. Glandular ridges absent; abdominal
separable from Isolda (cf. Day 1964).
notopodial rudiments present. Twelve thoracic un-
Pabits Chamberlin 1919c, P. deroderus Chamberlin cinigers. Anus surrounded by a circle of papillae.
1919c; only species. Pterampharete Augener 1918, P. luderitzi Augener
1918; only species.
paleae absent. Glandular ridges absent; notopodial AMPHARETINAE. Four pairs of pennate branchiae;
rudiments present in abdomen. Twelve thoracic un-
paleae present. Segment 4 without notosetae. Glan-
cinigers. Oral lobe prolonged.
dular ridges and abdominal notopodial rudiments ab-
Paiwa Chamberlin 1919c, P. abyssi Chamberlin 1919c; sent. Eleven thoracic uncinigers. Buccal tentacles
only species. papillose.
AMPHARETINAE. Four pairs of smooth branchiae; Pterolysippe Augener 1918, P. bipennata Augener
small paleae present. Glandular ridges absent; small 1918; only species.
abdominal notopodial rudiments present. Fourteen AMPHARETINAE. Four pairs of branchiae; three
thoracic uncinigers. smooth and one pennate; small paleae present. Glan-
Paramage Caullery 1944, P. madurensis Caullery dular ridges absent. Thirteen thoracic uncinigers.
1944; only species. Sabellides Milne Edwards in Malmgren 1966, Sabella
AMPHARETINAE. Four pairs of smooth branchiae. octocirrata Sars 1835; 8 species.
Paleae absent, first notosetae on segment 6; notopodial AMPHARETINAE. Four pairs of smooth branchiae;
lobes present on segments 4 and 5. Glandular ridges small paleae reported in some species. Glandular ridges
and abdominal notopodial rudiments absent. Eleven and abdominal notopodial rudiments absent. Eleven
thoracic uncinigers. thoracic uncinigers. Notosetae absent in segment 4.
Paramphicteis Caullery 1944, Sabellides angustifolia Buccal tentacles papillose.
Grube 1878; only species. Samytha Malmgren 1866, Sabellides sexcirrata Sars
AMPHARETINAE. Four pairs of smooth branchiae; 1856; 7 species.
paleae absent. Glandular ridges present; small ab- AMPHARETINAE. Three pairs of smooth branchiae;
dominal notopodial rudiments present. Fourteen tho- paleae absent. Glandular ridges absent; abdominal noto-
racic uncinigers. podial rudiments present. Fourteen thoracic uncinigers.
Parhypania Annenkova 1928, Amphicteis brevispinus Samythella Verrill 1873a, S. elongata Verrill 1873a;
Grube 1860; only species. 6 species.

AMPHARETINAE. Three pairs of smooth or flanged podial rudiments present; 12 thoracic uncinigers. Noto-
branchiae. Paleae absent. Glandular ridges absent; setae of last thoracic setiger modified.
abdominal notopodial rudiments absent. Twelve tho-
Weddellia Hartman 1967, W. profunda Hartman 1967;
racic uncinigers. Circle of anal papillae.
only species.
Samythopsis McIntosh 1885, S. grubei McIntosh 1885; AMPHARETINAE. Three pairs of smooth branchiae
only species. on three successive segments. Segment 3 with cap-
AMPHARETINAE. Three pairs of smooth branchiae; illary setae, but without paleal modifications. Glandular
paleae absent. Glandular ridges and abdominal noto- ridges absent; abdominal parapodia with long dorsal
podial rudiments present. Fourteen thoracic uncinigers. cirri. Fifteen thoracic uncinigers.
Schistocomus Chamberlin 1919a, S. hiltoni Chamberlin
1919a; 3 species.
AMPHARETINAE. Four pairs of branchiae, one Invalid Genera
smooth and three lamellate; paleae absent. Glandular Aryandes Kinberg 1867b, indeterminable
ridges absent; abdominal notopodial rudiments present. Branchiosabella Claparede 1863, see Ampharete
Twelve thoracic uncinigers. Two pairs of anal cirri. Crossostoma Gosse 1855, see Amphicteis
Sosane Malmgren 1866, S. sulcata Malmgren 1866; Eusamytha Hartman 1967, see Eusamythella
5 species. Heterobranchus Wagner 1885, see Sabellides
AMPHARETINAE. Four pairs of smooth branchiae; Melinnides Wesenberg-Lund 1950, see Melinnexis
paleae present. Glandular ridges absent; abdominal Annenkova, 1931
notopodial rudiments present. Twelve thoracic uncini- Melinnopsis Day 1955, see Melinnopsides
gers. Third from last thoracic notopodia elevated and Microsamytha Augener 1928a, see Alkmaria
with modified setae. Pseudosabellides Berkeley and Berkeley 1943, see
Asabellides
Sosanella Hartman 1965, S. apalea Hartman 1965; Rytocephalus Quatrefages 1866, indeterminable
only species.
paleae absent. Glandular ridges and abdominal noto-
podial rudiments absent. Thirteen thoracic uncinigers. FAMILY TEREBELLIDAE MALMGREN 1867
Notopodia in setiger 13 (third from last thoracic noto- Body in two regions; anterior region with biramous
podium) elevated with hirsute setae. parapodia and posterior region with neuropodia only.
Sosanides Hartmann-Schroder 1965, S. glandularis Prostomium a simple fold. Branchiae, when present,
Hartmann-Schroder 1965; only species. include one to three pairs on the first segments, asso-
AMPHARETINAE. Four pairs of smooth branchiae; ciated distinctly with separate segments. Uncini usually
with a large main fang and a crest of smaller teeth.
paleae present. Abdominal notopodial rudiments absent.
Twelve thoracic uncinigers. Setiger 11 with modified Terebellids are among the most common shallow-
notosetae. water polychaetes and are found in all environments.
The usually numerous buccal tentacles cannot be fully
Sosanopsis Hessle 1917, S. wireni Hessle 1917; 2 retracted into the mouth. They usually are grooved
species. and used in selective deposit-feeding on the surface.
AMPHARETINAE. Four pairs of smooth branchiae; Other forms may stretch the buccal tentacles into the
paleae absent. Glandular ridges absent; abdominal noto- water and capture particles from the water.
Key to Genera
Ia. A large scoop-shaped or conical proboscis present ARTACAMINAE 4

lb. Proboscis absent 2
2a (lb). Thoracic uncini in double or alternating rows in at least some setigers .... AMPHITRITINAE ... 14
2b (Ib). Thoracic uncini in single rows in all setigers 3
3a (2b). Branchiae present THELEPINAE 5
3b (2b). Branchiae absent POLYCIRRINAE 10
4a (la). Proboscis papillated, 17 thoracic setigers Artacama
4b (Iii). Proboscis grooved; 15 thoracic setigers Artacamella
5a (3a). First notosetae on first postbranchial segment Pseudampharete

5b (3a). First notosetae on one of the branchial segments 6

6a (5b). First notosetae on the first branchial segment Streblosoma
6b (5b). First notosetae on the second branchial segment 7
7a (6b). Lateral lappets present on one or a few anterior segments 8
7b (6b). Lateral lappets absent 9
8a (7a). Uncini from the fifth segment Euthelepus
8b (7a). Uncini from the first abdominal segment Telothelepus
9a (7b). Uncini from setiger 9 Parathelepus
9b (7b). Uncini from setiger 2 Thelepus
l0a (3b). Setae completely absent Hauchiella
l0b (3b). At least some setae present 11
lla(10b). Neurosetae absent 12
llb (l0b). Neurosetae present 13
12a (I I a). Thoracic notopodia vascularized and in part furcate or branched Enoplobranchus
I2b (11 a). Thoracic notopodia may be vascularized, but never furcate or branched Lysilla
13a (1lb). Neurosetae short-handled uncini Polycirrus
13b (1lb). Neurosetae long-handled spines Amaeana
14a (2a). Branchiae absent 15
1 4b (2a). Branchiae present 22
15a (14a). Seventeen or fewer thoracic setigers 16
15b (14a). More than 20 thoracic setigers 21
16a (l5a). All notosetae smooth-tipped 17
16b (15a). At least some notosetae with denticulated tips 19
17a (16a). Lateral lappets absent; 17 thoracic setigers Laphania
I7b (16a). Lateral lappets present on one or a few anterior segments; 16 thoracic setigers 18
I8a (17b). Third segment with transverse ridge across dorsum Leaena
I8b (17b). Third segment without transverse ridge Stschapovella
19a (16b). Neurosetae from setiger 3; some notosetae smooth Proclea
I 9b (16b). Neurosetae from setiger 2; all notosetae denticulated 20
20a (19b). Fifteen thoracic setigers; notosetae finely denticulated Lanassa
20b (19b). Fourteen thoracic setigers; some notosetae distinctly pectinate Phisidia
21a (15b). All notosetae tapering, distally smooth-tipped Baffinia
21b (15b). Some notosetae subdistally inflated, distally denticulated Spinosphaera
22a (14b). Notosetae with marginally serrated tips 23
22b (14b). Notosetae with smooth tips 31
23a (22a). Three anterior segments with large lateral lappets to which the branchiae are attached . Colymmatops
23b (22a). Lateral lappets present or absent, but never directly associated with the branchiae 24
24a (23b). Uncini from third thoracic setiger Neoleprea
24b (23b). Uncini from second thoracic setiger 25
25a (24b). Sixteen or 17 thoracic setigers 26
25b (24b). Eighteen or more thoracic setigers 28
26a (25a). A single pair of branchiae; 16 thoracic setigers Spiroverma
26b (25a). Two or three pairs of branchiae; 17 thoracic setigers 27
27a (26b). Lateral lappets present; branchiae stalked and branched Neoamphitrite
27b (26b). Lateral lappet absent; branchiae sessile filaments Amphitrite
28a (24b). Branchiae attached on segments 2-4 29
28b (24b). Branchiae attached on another combination of segments 30
29a (28a). Branchiae arborescent from the base Terebella
29b (28a). Branchiae distinctly stalked Amphitritides
30a (28b). Branchiae on segments 1, 2 and 5 Polymniella
30b (28b). Branchiae on segments 3, 7 and 13 Terebellobranchia
31a (22b). Anterior uncini long-handled 32
31b (22b). All uncini short-handled 35
32a (31 a). Notosetae from segment 5, uncini from segment 6 Opisthopista
32b (31 a). Notosetae from segment 4, uncini from segment 5 33
33a (32b). Branchiae smooth, ampharetinlike Eupistella

33b (32b). Branchiae branched 34

34a (33b). Lateral lappets at least on segments 2 and 4 Pista
34b (33b). Lateral lappets limited to segment 3 Lanicides
35a (31b). Ventrum with large anteriorly opening glandular folds on segments 3-5 Pararionice
35b (31b). Ventrum with glandular scutes or smooth 36
36a (35b). Lateral lappets present on one or a few anterior segments 37
36b (35b). Lateral lappets inconspicuous or absent 44
37a (36a). Sixteen thoracic setigers; branchiae very long-shafted Axionice
37b (36a). Seventeen or 18 thoracic setigers; branchiae short-shafted or sessile 38
38a (37b). Eighteen thoracic setigers Melinella
38b (37b). Seventeen thoracic setigers 39
39a (38b). First four segments flattened with large lateral lappets forming an anterior plaque; one pair of
branchiae Scionella
39b (38b). First four segments not modified into a flattened plaque; lateral lappets varying in size; two or three
pairs of branchiae 40
40a (39b). Branchiae tufts of sessile filaments Thelepides
40b (39b). Branchiae branching off one or a few stems 41
41a (40b). Buccal lateral lappets form a crest across the dorsum Paralanice
4lb (40b). No dorsal crest across the buccal segment 42
42a (41b). Uncini with teeth in a single row Loimia
42b (41 b). Uncini with teeth in two or more rows 43
43a (42b). Small lateral lappets on segments 2 and 3; tube opening unadorned Eupolymnia
43b (42b). Large lateral lappets on segments 2 and 4; tube opening with fan-shaped frills Lanice
44a (36b). Thirteen thoracic setigers Ramex
44b (36b). At least 15 thoracic setigers 45
45a (44b). First notosetae on first branchial segment Naneva
45b (44b). First notosetae on last branchial segment or from one of the first postbranchial segments 46
46a (45b). Uncini present from first setiger Reteterebella
46b (45b). Uncini present from second setiger 47
47a (46b). Uncini in an open circle in posterior thoracal setigers Terebellanice
47b (46b). Uncini arranged back to back in posterior thoracal setigers 48
48a (47b). Two pairs of branchiae; thorax with 15 to 40 setigers Nicolea
48b (47b). Three pairs of branchiae; thorax with 17 setigers Scionides
Generic Definitions Artacama Malmgren 1866, A. proboscidea Malmgren

Amaeana Hartman 1959, Polycirrus trilobatus Sars 1866; 8 species.
1863; 5 species. ARTACAMINAE. Three pairs of filamentous
POLYCIRRINAE. Tentacular lobe trifoil-shaped. branchiae on segments 2-4. Lateral lappets absent.
Ten to 13 thoracic segments; notosetae from segment 3. Seventeen thoracic setigers; notosetae distally smooth.
Uncini absent; abdominal neurosetae long-shafted Papillose proboscis attached ventrally on the peris-
spines. tomium.
Amphitrite O.F. Muller 1771, A. cirrata O.F. Muller Artacamella Hartman 1955, A. hancocki Hartman
AMPHITRITINAE. Eyes rarely present. Three pairs ARTACAMINAE. Three pairs of smooth, am-
of sessile branchiae from segment 2; nephridial papillae pharetinlike branchiae on segments 2-4. Fifteen tho-
on segment 3 and from segment 6. Lateral lappets racic setigers; uncini in all thoracic setigers; each
absent. Notosetae from fourth segment; distally ser- uncinus long-shafted. Grooved, boat-shaped proboscis
rated; 13 to 25 thoracic setigers. attached ventrally on the peristomium.
Amphitritides Augener 1922, Terebella gracilis Grube Axionice Malmgren 1866, Terebella fexuosa Grube
AMPHITRITINAE. Two pairs of branching, stalked AMPHITRITINAE. Two or three pairs of long-
branchiae from segment 2. Lateral lappets absent. shafted, branched branchiae. Sixteen thoracic setigers;
Serrated notosetae present from segment 4; uncini face lateral lappets present. Notosetae distally smooth; all
to face in posterior thoracic setigers. uncini short-handled.

Baffinia Wesenberg-Lund 1950, B. multisetosa Wesen- Lanice Malmgren 1866, Nereis conchilega Pallas
berg-Lund 1950; only species. 1766; 8 species.
AMPHITRITINAE. Branchiae absent. Notosetae AMPHITRITINAE. Three pairs of branched bran-
present from third segment to the end of the body chiae. Lateral lappets present. Seventeen thoracic seti-
(more than 70 segments); uncini present from second gers. Smooth-tipped notosetae from segment 4; uncini
setiger; uniserial in first eight uncinigers, then biserial back to back in posterior thoracic setigers. Tube with
and finally uniserial in last 30-35 setigers. Capillary branched fine-meshed fan attached to opening.
setae distally smooth.
Lanicides Hessle 1917, Terebella (Phyzelia) bilobata
Bathya Saint-Joseph 1894, Leaena abyssorum Mc- Grube 1877; 3 species.
Intosh 1885; 3 species. AMPHITRITINAE. Two pairs of branched bran-
AMPHITRITINAE. Branchiae absent. Uncini with chiae; lateral lappets present. Smooth-tipped notosetae
short handles; crested; capillary distally smooth, represent from segment 4; long-shafted uncini present
sembles Proclea in setal structures. Incompletely from segment 5.
described.
Laphania Malmgren 1866, L. boecki Malmgren 1866;
Colymmatops Peters 1854, C. granulatus Peters 1854; only species.
only species. AMPHITRITINAE. Branchiae absent. Lateral lap-
AMPHITRITINAE. Three first segments with large pets absent. Seventeen thoracic setigers; notosetae
lateral lappets to which are attached branchiae. Thirteen distally smooth. Uncini present from setiger 7.
or 14 thoracic setigers. Notosetae distally serrated.
Incompletely described. Leaena Malmgren 1866, Terebella abranchiata Sars
1865; 10 species.
Enoplobranchus Webster 1879, Chaetobranchus san- AMPHITRITINAE. Branchiae absent. Lateral lap-
guinea Verrill 1873b; only species. pets present; third segment with transverse ridge across
POLYCIRRINAE. Notopodial lobes prolonged, vas- dorsum. Smooth-tipped notosetae present from seg-
cularized and in part furcate or branched. Uncini absent; ment 4. Sixteen thoracic setigers.
notosetae spinose capillaries, with usually one seta
much longer than the others. Loimia Malmgren 1866, Terebella medusa Savigny
1818; 16 species.
Eupistella Chamberlin 1919c, Eupista darwini Mc-
AMPHITRITINAE. Three pairs of branched bran-
Intosh 1885; 4 species.
chiae. Lateral lappets present. Seventeen thoracic seti-
AMPHITRITINAE. Two pairs of smooth, am-
gers; notosetae smooth-tipped. Uncini with all teeth in
pharetinlike branchiae. Seventeen thoracic setigers;
a single row (pectinate).
some anterior uncini with prolonged shafts, notosetae
distally smooth. Lysilla Malmgren 1866, L. loveni Malmgren 1866;
Eupolymnia Verrill 1900, Amphitrite nesidensis delle 10 species.
Chiaje 1828; 12 species. POLYCIRRINAE. Six to 12 thoracic segments; noto-
AMPHITRITINAE. Three pairs of branching bran- setae from segment 3. Neurosetae completely absent.
chiae; lateral lappets on segments 2-3. Smooth-tipped Melinella McIntosh 1914, M. macduffi McIntosh 1914;
notosetae from segment 4; 17 thoracic setigers. only species.
Euthelepus McIntosh 1885, E. setubalensis McIntosh AMPHITRITINAE. One pair of branched branchiae;
1885; 6 species. 18 thoracic setigers, all with uncini. Lateral lappets
THELEPINAE. Branchiae on segments 2-4, some- present.
times as single filaments only. Lateral lappets present. Naneva Chamberlin 1919a, N. hespera Chamberlin
Notosetae from second branchial segment; present on 1919a; only species.
20 segments. Uncini first present from first post- AMPHITRITINAE. Two pairs of dendritically
branchial segment. branched branchiae from segment 2; smooth-tipped
Hauchiella Levinsen 1893, Polycirrus tribullata Mc- notosetae present from first branchial segment. Twenty-
Intosh 1869; only species. seven thoracic setigers. Lateral lappets absent. Uncini
POLYCIRRINAE. Thorax of ten segments; usually present in double-rows in most thoracic segments.
about 70 segments in all. All setae absent.
Neoamphitrite Hessle 1917, Amphitrite affinis Malm-
Lanassa Malmgren 1866, L. nordenskioldi Malmgren gren 1866; 11 species.
1866; 7 species. AMPHITRITINAE. Three pairs of branched bran-
AMPHITRITINAE. Branchiae absent. Lateral lap- chiae; lateral lappets present. Seventeen thoracic seti-
pets sometimes present. Fifteen thoracic setigers. Noto- gers with distally serrated notosetae. Nephridial papillae
setae present from segment 4, denticulated tips. present from segment 3.

Neoleprea Hessle 1917, Leprea streptochaeta Ehlers Polycirrus Grube 1850, P. medusa Grube 1850; 39
1897; 5 species. species.
AMPHITRTTINAE. Two or three pairs of branched POLYCIRRINAE. Thorax with a variable number
branchiae; lateral lappets absent. Notosetae first pres- of setigers; notosetae present from segment 3. Uncini
ent from segment 3; some smooth, some distally den- first present from segments 7-18. Notosetae distally
ticulated. Seventeen-40 thoracic setigers. Nephridial smooth or serrated.
papillae present on segments 3-9.
Polymniella Verrill 1900, P. aurantiaca Verrill 1900;
Nicolea Malmgren 1866, Terebella zostericola Orsted only species.
1844; 22 species. AMPHITRITINAE. Three pairs of branched bran-
AMPHITRITINAE. Two pairs of branched branchiae. chiae on segments 1, 2 and 5. Notosetae from segment
Lateral lappets absent. Smooth-tipped notosetae present 1, marginally dentate; uncini from segment 2. Twenty-
from segment 4; 15-40 thoracic setigers. Uncini ar- two or more thoracic segments.
ranged back to back in posterior thoracic segments.
Proclea Saint-Joseph 1894, Leaena graffli Langerhans
Opisthopista Caullery 1944, O. sibogae Caullery 1944; 1880; 3 species.
only species. AMPHITRITINAE. Branchiae absent. Lateral lap-
AMPHITRTTINAE. Two pairs of branched branchiae; pets present. Sixteen thoracic setigers; notosetae distally
lateral lappets present on at least segments 2 and 4. First either dentate or smooth. Uncini from setiger 3.
notosetae in segment 5 and first uncini in segment 6.
Pseudampharete Hartmann-Schrdder 1960b, P. ten-
Anterior uncini long-shafted.
taculata Hartmann-Schroder 1960b; only species.
Paralanice Caullery 1944, P. timorensis Caullery 1944; THELEPINAE. Two pairs of sessile branchial fila-
only species. ments on large bosses on first and second segment.
AMPHITRTTINAE. Three pairs of branched bran- Notosetae first present from first postbranchial seg-
chiae; large lateral buccal lappets connected across ment; uncini from setiger 7.
dorsum with a crest; lateral lappets also on segments
Ramex Hartman 1944b, R. californiensis Hartman
2 and 3. Seventeen thoracic setigers; smooth-tipped
1944b; only species.
capillaries from segment 4.
AMPHITRTTINAE. One pair of branched branchiae
Parathelepus Caullery 1915, Thelepides collaris South- on second segment. Notosetae from segment 4; thorax
ern 1914; only species. with 13 setigers; notosetae distally smooth.
THELEPINAE. Three pairs of branchiae; lateral Reteterebella Hartman 1963b, R. queenslandia Hart-
lappets absent. Notosetae from second branchial seg- man 1963b; only species.
ment; uncini from setiger 9. AMPHITRITINAE. Three pairs of branched bran-
Pararionice Fauchald 1972, P. artifex Fauchald 1972; chiae; lateral lappets inconspicuous. Sixteen thoracic
only species. setigers; notosetae from segment 4; distally smooth.
AMPHITRITINAE. One pair of branched branchiae Uncini present from first setiger.
with double bases on segments 3 and 4. Sixteen tho- Scionella Moore 1903, S. japonica Moore 1903; 4
racic setigers; notosetae distally smooth. Ventral part species.
of segments 3-5 covered by a large glandular apparatus AMPHITRITINAE. One pair of branchiae on seg-
that opens anteriorly on segment 3 in two trumpet- ment 4. Seventeen thoracic setigers; notosetae first
shaped openings. present on segment 4; notosetae distally smooth. Four
Phisidia Saint-Joseph 1894, Leaena oculata Langerhans first segments flattened dorsoventrally with very large
1880; 3 species. longitudinally oriented lateral lappets, forming a large
AMPHTTRTTINAE. Branchiae absent; lateral lappets oblique plaque at the anterior end.
absent. Notosetae distally denticulate; longer finely so, Scionides Chamberlin 1919b, Terebella reticulata
the shorter coarse, with pectinate appearance. Uncini Ehlers 1887; 2 species.
from setiger 2; 14 thoracic setigers. AMPHITRITINAE. Three pairs of branched bran-
Pista Malmgren 1866, Amphitrite cristata O.F. Muller chiae; seventeen thoracic setigers; notosetae from seg-
ment 4; notosetae distally smooth. Uncini arranged back
1776; 40 species.
to back in posterior thoracic segments.
AMPHITRTTINAE. Two pairs of stalked branched
branchiae; lateral lappets large, on segments 2 and 4 Spinosphaera Hessle 1917, S. pacifica Hessle 1917;
at least. Smooth-tipped notosetae from segment 4; 2 species.
long-handled uncini present in anterior setigers. Fifteen AMPHTTRTTINAE. Branchiae absent. Twenty-three
to 24 thoracic setigers. or more thoracic setigers; uncini present from setiger 2.
Notosetae denticulate; the longer ones with hispid Thelepus Leuckart 1849, Amphitritee cincinnata Fabri-
swellings subdistally. Lateral lappets absent. cius 1780; 32 species.
THELEPINAE. Sessile, filiform branchiae on seg
Spiroverma Uchida 1968, S. ononokomachii Uchida ments 2-4; smooth-tipped notosetae present from
1968; only species.
second branchial segment (segment 3). Uncini from
AMPHITRTTINAE. One pair of sessile branchiae,
segment 5.
each with maximally eight filaments on segment 2.
Sixteen thoracic setigers; notosetae marginally serrate.
Body strongly spiralled.
Taxonomic Notes
Streblosoma Sars 1872, Grymaea bairdi Malmgren
Day (1967) altered the definition of the sub-families
1866; 20 species.
to include the abranchiate members of AMPHITRI-
THELEPINAE. Three pairs of sessile branchiae on
TINAE among the POLYCIRRINAE. This change
segments 2-4 (may be absent). Notosetae from first
appears unfortunate, in that these genera resemble
branchial segment (segment 2). Uncini from seg-
branchiate members of the AMPHITRTTINAE very
ment 5.
closely in setal structures as well as in the structure of
Stschapovella Levenstein 1957, S. tatjanae Leven- the anterior end. The treatment here reflects this view.
stein 1957; only species. The genus Pseudampharete has been included among
AMPHITRITINAE. Branchiae absent; lateral lap- the THELEPINAE since it is branchiate and has the
pets present. Smooth-tipped notosetae from segment 4; uncini in single rows in all thoracic segments. It further
uncini from segment 5. Sixteen thoracic setigers. resembles members of this subfamily in that the branch-
Notosetae finely capillary rather than limbate. All iae are sessile filaments. However, this latter feature
nephridia free from one another. may also be present among members of the AMPHITRI-
TINAE. The placement must be considered temporary.
Telothelepus Day 1955, T. capensis Day 1955; only
The genus Bathya Saint-Joseph 1894, listed above
species.
in the definitions has not been included in the key. It
THELEPINAE. Two or three pairs of branchiae from
belongs in the abranchiate gorup of AMPHITRITINAE,
segment 2. Notosetae from second branchial segment
and is very poorly known.
(segment 3). Lateral lappets present. Fifteen thoracic
The genus Pseudothelepus Augener 1918 is con-
setigers. Tentacular lobe very large and frilly. Neuro-
sidered here a synonym of Streblosoma, as suggested
setae absent on thorax, uncini present on abdomen.
by Day (1967).
Terebella Linnaeus 1767, T. lapidaria Linnaeus 1767; Some of the genera are difficult to separate from re-
28 species. lated forms; no revision was attempted on this occasion.
AMPHITRTTINAE. Two or three pairs of branched
branchiae; lateral lappets absent. Thorax with variable,
usually large, number of setigers; notosetae from seg- Invalid Genera
ment 4; distally serrated. Uncini face to face in pos-
Amaea Malmgren 1866, see Amaeana
terior thoracic segments.
Amphiro Montagu 1808, see Amphitrite
Terebellanice Hartmann-Schri der 1962b, T. laeviseta Amphitritoides Costa 1862, see Eupolymnia
Hartmann-Schri der 1962b; only species. Amphytrite Renier 1804, indeterminable
AMPHITRITINAE. Two pairs of branchiae from Anisocirrus Gravier 1905a, see Polycinrus
third segment; lateral lappets absent. Notosetae smooth- Aphlebina Claparede 1864, see Polycirrus
tipped. Uncini in an open circle on posterior thoracic Apneuma Quatrefages 1866, see Polycirrus
segments. Athelepus Chamberlin 1919c, NoMsnNUDUM
Chaetobranchus Verrill 1873b, see Enoplobranchus
Terebellobranchia Day 1951, T. natalensis Day 1951;
Cyaxares Kinberg 1867b, see Polycirrw
2 species.
Dejoces Kinberg 1867b, see Polycirrus
AMPHITRTTINAE. Three pairs of branched bran-
Dendrobranchus Wagner 1885, indeterminable
chiae on segments 3, 7 and 13. Distally serrated noto- Dendrophora Grube 1870a, see Pista
setae present from segment 4; more than 19 thoracic
Ehlersiella McIntosh 1885, indeterminable
setigers present. Ereutho Malmgren 1866, see Polycirrus
Thelepides Gravier 191 la, T. koehleri Gravier 1911 a; Eugrymaea Verrill 1900, see Streblosoma
3 species. Eupista McIntosh 1885, see Eupistella
AMPHITRTTINAE. Three pairs of filiform branchiae; Euscione Chamberlin 1919c, see Axionice
lateral lappets present. Smooth-tipped notosetae from Grymaea Malmgren 1866, see Streblosoma
segment 3; 17 thoracic setigers present. Heterophenacia Quatrefages 1866, see Thelepus

Heterophyselia Quatrefages 1866, see Terebella Uncinochaeta Quatrefages 1866, indeterminable

Heteroterebella Quatrefages 1866, see Terebella Venusia Johnston 1865, see Thelepus
Idalia Quatrefages 1866, see Pista Wartelia Giard 1878, see Lanice
Laphaniella Maim 1874, see Lanassa
Leprea Malmgren 1866, see Terebella
Leucariste Malmgren 1866, see Polycirrus FAMILY TRICHOBRANCHIDAE MALMGREN 1 866
Lumara Stimpson 1854, see Thelepus Body separated into two regions; anterior region with
Neottis Malmgren 1866, see Thelepus biramous parapodia, posterior region with neuropodia
Odysseus Kinberg 1867b, indeterminable only. Prostomium a large hood over the mouth. Max-
Otanes Kinberg 1867b, see Pista i mally four pairs of branchiae present on anterior
Pallonia Costa 1862, see Eupolymnia setigers. Anterior neuropodia with long-shafted hooks;
Phenacia Quatrefages 1866, see Thelepus abdomen with densely crested uncini. Notosetae
Pherea Saint-Joseph 1894, see Lanassa capillary.
Phyzelia Schmarda 1861, see Nicolea The trichobranchids are related closely to the tere-
Polymnia Malmgren 1867, see Eupolymnia bellids and frequently are included among them as a
Protothelepus Verrill 1900, see Euthelepus subfamily (Day 1967). They differ rather strikingly
Schmardanella McIntosh 1885, see Terebella in body-proportions, in that they tend to be slenderer
Scione Malmgren 1866, see Axionice and more muscular than the terebellids. The large
Scionopsis Verrill 1873b, see Pista folded prostomium also gives them a characteristic
Solowetia Ssolowiew 1899, see Proclea appearance. One species, Terebellides stroemi, has
Thelepella Chamberlin 1919c, see Nicolea been reported from worldwide, areas, and is apparently
Thelephusa Verrill 1871, see Thelepus nearly everywhere, an important member of soft-bottom
Thelepides Southern 1914, see Parathelepus environments. Members of the family appear to be
Thelepodopsis Sars 1872, see Thelepus most common in cold-water soft bottoms and may be
Torquea Leidy 1855, see Polycirrus represented best in deep water.
Key to Genera
la. A single middorsally attached branchia present 2
lb. At least a pair of branchiae present 3
2a (la). Branchia a single tapering, digitate projection Unobranchus
2b (Ia). Branchia stalkedd with four lobes; each lobe with numerous flat branchial lamellae Terebellides
3a (lb). Two or three pairs of branchiae present 4
3b (lb). Four pairs of branchiae present 5
4a (3a). Two pairs of branchiae, 17 thoracic setigers Filibranchus
4b (3a). Three pairs of branchiae; 15 thoracic setigers Trichobranchus
5a (3b). Each branchia with pectinate branchial lamellae Ampharetides
5b (3b). Each branchia simple and digitate or rosette-shaped 6
6a (5b). All branchiae simple and tapering Octobranchus
6b (5b). Fourth pair of branchiae sessile rosettes Novobranchus
Generic Definitions Novobranchus Berkeleyy and Berkeley 1954, N. pacificus

Ampharetides Ehlers 1913, A. vanhoeffeni Ehlers 1913; Berkeley and Berkeley 1954; only species.
only species. Sixteen thoracic setigers; uncini present from setiger
Seventeen thoracic setigers; uncini'present on nine 4. Four pairs of branchiae on segments 2-5; first three
last thoracic setigers and on abdomen. Four pairs of pairs lanceolate; last pair a frilled rosette.
branchiae on first setiger; each with pectinate branchial Octobranchus Marion and Bobretzky 1875, Terebella
lamellae. lingulata Grube 1863; 4 species.
Filibranchus Mahn 1874, F. roseus Malm 1874; only Sixteen thoracic setigers; first setiger is segment 3;
species. first uncini on setiger 4. Four pairs of similar lanceolate
Fifteen thoracic setigers; all with uncini. Two pairs branchiae.
of smooth branchiae on the second and third segment Terebellides Sars_ 1835, T. stroemi Sars 1835; 14
(presetal branchiae). species.

/976 THE POLYCHAETE WORMS 13 5
Eighteen thoracic setigers; first setiger is segment 3; Invalid Genera

uncini present from setiger 6. Single dorsal branchia Aponobranchus Gravier 1905a, see Terebellides
with four lamellate branchial lobes. Corephorus Grube 1846, see Terebellides
FAMILY BOGUEIDAE HARTMAN AND

Trichobranchus Malmgren 1866, T. glacialis Malm-
gren 1866; 5 species. FAUCHALD 1971
Fifteen thoracic setigers; first setae on segment 6; Body slender and cylindrical with few segments;
all thoracic setigers with uncini. Three pairs of simple, prostomium rounded without appendages. Parapodia
lanceolate branchiae. biramous with long slender, in part spinose notosetae
and short-handled, avicular crested uncini in the neuro-
Unobranchus Hartman 1965, U. abyssalis Hartman podia, in part in double rows.
1965; only species. The bogueids, known for two genera, have setae
Twelve thoracic setigers; first setae on segment 3; resembling the terebellid uncini, but lack completely
uncini first present on setiger 4. A single, large lance- anterior appendages. They are known from a few loca-
olate branchia present mid-dorsally on segment 2. tions in the western Atlantic Ocean only.
Key to Genera
First three setigers without neurosetae Boguea

First four setigers without neurosetae Boguella
Generic Definitions are strictly sessile and never leave their tubes; the
smaller forms, such as species of Fabricia and allied
Boguea Hartman 1945, B. enigmatica Hartman 1948; genera, are capable of moving around. Most of the
only species. l arger forms are associated with shallow water; smaller
Two anterior asetigerous segments; three first species are common in deep sea collections.
setigers without neurosetae. The major characteristics used to identify the sabel-
lids, include the presence or absence of companion
Boguella Hartman and Fauchald 1971, B. ornata Hart-
setae to the neuropodial uncini in the thorax; these
man and Fauchald 1971; only species.
also have been called pennoned setae or pick-ax setae.
One anterior asetigerous segment; four first setigers
The neutral term companion seta is preferred here.
without neurosetae.
They occur in an anterior row, in front of the uncini
they accompany and usually are small and deeply im-
ORDER SABELLIDA bedded in the epidermal tissues. They are more easily
seen by the reflection they give off under a stereo
Prostomium reduced, fused with the peristomium microscope, than in microscopic preparation under
which usually forms a large tentacular crown; setae the compound microscope. The structure of the ten-
include thoracic notopodial limbate and geniculate tacular crown such as the number of radioli, and the
kinds and neuropodial uncini; setal positions reversed presence of small, external appendages called stylodes,
in the abdomen. are important.
One of the key features in the group lies in the struc-
ture of the thoracic uncini. These may be acicular, by
FAMILY SABELLIDAE MALMGREN 1867 which is meant that the crested head, with one large
tooth and several smaller ones, is supported by a gently
Body cylindrical with a thorax of few setigers and curved, often nearly straight shaft. By avicular uncini
abdomen with few to many. Uncini crested or with teeth are meant uncini that are essentially Z-shaped (some-
in several rows; long- or short-handled. Tube present times called swan-shaped) with the small crested head
in most species, made of varying material, but never at the top of the Z, and the shaft sharply bent. The
calcareous. shafts of these Z-shaped uncini may be short or long.
Sabellids characteristically have nearly smooth ap- In the key below, attempts have been made to avoid
pearing bodies, cylindrical and tapering posteriorly, the more confusing part of the terminology, but a
with large, often maroon or red-colored tentacular complete avoidance of this terminology has not been
crowns. Most of the forms, especially the larger ones, possible.

FIGuRE 37. (A), Family SABELLIDAE, Chone sp., off Santa Catalina Island, 50 m, 18x; (B), Family SABELLONGIDAE,
Sabellonga disjuncta, anterior end in right ventrolateral view, modified after Hartman, 1969, 33x; (C), posterior end of the
above, 33x; (D), Family CAOBANGIIDAE, Caobangia abbotti, after Jones, 1974, about 17x; (E), right lateral view of the above,
about 17x.
Key to Genera
Ia. Abdominal uncini form nearly complete cinctures around the body MYXICOLINAE
Myxicola
lb. Abdominal uncini in short, discrete tori 2
2a (Ib). Thoracic uncini with long, gently curved shafts (acicular) companion setae always absent
. . . . . . . . . . . . . . . . ............................................FABRICINAE 3
2b (lb). Thoracic uncini with short or long, but always strongly bent shafts. (avicular); companion setae
present in some forms SABELLINAE 15
3a (2a). Abdomen with two or three setigers 4
3b (2a). Abdomen with four or more setigers 9

4a (3a). Radioles partially united by a membrane Monroika

4b (3a). Radioles free to the base 5
5a (4b). With an extensive lower lip, curved up under the weakly bilobed prostomium Pseudofabricia
5b (4b). Without a distinct lower lip; prostomium anteriorly rounded 6
6a (5b). Abdominal uncini short-handled Fabriciola
6b (5b). Abdominal uncini long-handled 7
7a (6b). Without ventral vascularized filaments Fabricia
7b (6b). With ventral vascularized filaments in the tentacular crown 8
8a (7b). Ventral vascularized filaments branched; three pairs of radioli Augeneriella
8b (7b). Ventral vascularized filaments smooth; two pairs of radioli Manayunkia
9a (3b). Posterior end distinctly modified, with fused segments or greatly expanded pygidium 10
9b (3b). Posterior end not modified 11
l0a (9a). Seven last setigers covered dorsally by the large, ring-shaped pygidium Euchonella
l0b (9a). Last several setigers flanged laterally to form a spoon-shaped dorsal cavity Euchone
IIa(9b). Radioles united by a membrane Chone
1lb (9b). Radioles free to the base 12
12a (l Ib). Abdominal uncini with a large main fang and a crest of smaller teeth Jasmineira
12b (l lb). Abdominal uncini with major teeth in two or more rows 13
13a (12b). More than ten pairs of radioli in the tentacular crown Dialychone
13b (12b). Maximally five pairs of radioli in the tentacular crown 14
14a (13b). Radioles externally distinctly ridged Oriopsis
14b (13b). Radioles externally rounded Desdemona
15a (2b). Thoracic neuropodial companion setae absent 16
15b (2b). Thoracic neuropodial companion setae present 21
1 6a (15a). Radioli with external stylodes 17
1 6b (15a). Radioles without external stylodes 18
17a (16a). Four thoracic setigers, external stylodes very small Pseudobranchiomma
17b (16a). Eight thoracic setigers; external stylodes well developed Branchiomma
18a (16b). Thoracic spatulate notosetae present 19
18b (16b). Thoracic spatulate notosetae absent 20
19a (18a). Thoracic uncini with gently curved, long handles Fabrisabella
19b (18a). Thoracic uncini with sharply bent handles Laonome
20a (18b). Collar reduced, radioli basally united by a web Euratella
20b (18b). Collar well developed; radioli free to the base Sabellastarte
21a (15b). Tentacular crown with radioli arranged in one or two spirals 22
21b (15b). Tentacular crown with radioli in two semicircles 26
22a (21 a). Posterior thoracic notopodia with lancet-shaped, transversely striated setae Bispira
22b (21 a). Lancet-shaped notopodial setae absent 23
23a (22b). Collar two-lobed 24
23b (22b). Collar four-lobed 25
24a (23a). Anterior ventrum irregularly rugose Distylidia
24b (23a). Anterior ventrum smooth, apart from the glandular fields Demonan
25a (23b). One half of tentacular crown very much larger than the other; spatulate setae absent . . Spirographis
25b (23b). Both parts of the tentacular crown similar in size; spatulate setae present Eudistylia
26a (21b). Large subdistal compound eyes on a few dorsal radioli Megalomma
26b (21b). Eyes, if present, smaller and scattered over most of the radioli 27
27a (26b). Collar reduced or absent, a triangular ventral projection at base of tentacular crown present
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ..................Amphiglena
27b (26b). Collar well developed; triangular ventral projection absent 28
28a (27b). Notosetae of the collar-segment in a long row Hypsicomus
28b (27b). Notosetae of the collar-segment in a tuft 29
29a (28b). Radioli dichotomously branching Schizobranchia
29b (28b). Radioli not divided 30
30a (29b). Spatulate thoracic notosetae absent Sabella
30b (29b). Spatulate thoracic notosetae present 31
3 la (30b). Thoracic uncini at least in part nearly straight and long-handled 32

31b (30b). Thoracic uncini sharply bent, long- or short-handled 33

32a (31 a). Abdominal uncini long-handled and nearly straight Potaspina
32b (31 a). Abdominal uncini short-handled and strongly bent Potamethus
33a (31b). Thoracic uncini long-handled Panousea
33b (31b). Thoracic uncini short-handled Potamilla
Generic Definitions Desdemona Banse 1957, D. ornata Banse 1957; only

species.
Amphiglena Claparede 1864, Amphicora mediterranea FABRICINAE. Three pairs of radioli, radiolar backs
Leydig 1851; 3 species. rounded, webbing absent. Collar divided dorsally and
SABELLINAE. Radioles in semi-circles; external united ventrally. Abdominal uncini with short, quad-
stylodes, eyes and webbing absent. Collar absent; tri-
rangular bases and teeth in two rows along one side.
angular ventral projection present at base of tentacular Abdomen with four to 12 setigers.
crown. Approximately eight thoracic setigers present;
abdominal uncini strongly bent, crested. Dialychone Claparede 1870a, D. acustica Claparede
1870a; only species.
Augeneriella Banse 1957, A. hummelincki Banse 1958; FABRICINAE. Twelve or more pairs of radioli;
4 species. webbing absent. Collar well developed on both sides.
FABRICINAE. Three pairs of radioli, branchial Abdominal uncini with short, quadrangular base and
heart present; collar dorsally low, ventrally higher.
teeth in two or more rows along one side. Abdomen
Forked ventral vascularized filament present. Eight
with several setigers.
thoracic and three abdominal setigers; abdominal uncini
long-handled. Distylidia Hartman 1961, Distylia rugosa Moore 1904b;
2 species.
Bispira Kroyer 1856, Amphitrite volutacornis Montagu SABELLINAE. Radioles spiralled; eyes present,
1804; 3 species. external stylodes and webbing absent. Collar bilobed.
SABELLINAE. Radioles in spirals; eyes present, Both thoracic and abdominal uncini long-handled, but
stylodes and webbing absent. Collar two- or four-lobed.
sharply bent. Anterior ventrum irregularly rugose.
Number of thoracic segments variable. Spatulate tho-
racic notosetae absent; late thoracic notopodia with Euchone Malmgren 1866, Sabella analis Kroyer 1856;
lancent-shaped, transversely striated setae. 30 species.
FABRICINAE. Numbers of radioles variable; web-
Branchiomma Kolliker 1858, Amphitrite bombyx bing present. Collar usually well developed. Last
Dalyell 1853; 21 species. abdominal setigers flanged laterally to form a deep
SABELLINAE. Radioles sometimes spiralled, some- spoon-shaped cavity. Abdominal uncini avicular. Ab-
times in semicircles; eyes and external stylodes present, domen with numerous setigers.
webbing absent. Collar well developed. Thoracic
spatulate setae absent; thoracic neuropodial companion Euchonella Fauchald 1972, E. magna Fauchald 1972;
setae absent. Eight thoracic segments only species.
FABRICINAE. Twelve pairs of radioli; webbing
Chone Kroyer 1856, C. infundibuliformis Kroyer and eyes absent. Collar well developed, highest ven-
1856; 26 species. trally. Pygidium expanded to a large dorsal horse-shoe
FABRICINAE. More than five, usually many pairs covering the last seven setigers. Abdominal uncini
of radioli present; distinct webbing present between avicular. Abdomen with numerous setigers.
the radioli. Collar well developed. Abdominal uncini
with short, quadrangular base and crested tip. Ab- Eudistylia Bush 1904a, Sabella vancouveri Kinberg
domen with several setigers. 1867b; 3 species.
SABELLINAE. Radioles spiralled; eyes present,
Demonax Kinberg 1867b, D. leucaspis Kinberg 1867b; webbing and external stylodes absent. Collar four-
8 species. lobed. Spatulate thoracic notosetae present. Abdominal
SABELLINAE. Radioli in spiral; eyes present; uncini avicular.
webbing and external stylodes absent. Collar bilobed.
Spatulate notosetae and neuropodial companion setae Euratella Chamberlin 1919c, Laonome salmacidis
present in thorax. Both thoracic and abdominal uncini Claparede 1870a; 2 species.
with long handles, but very sharply and distinctly SABELLINAE. Radioli in semi-circles; webbing and
bent and distally crested. eyes present; external stylodes absent. Collar reduced.

Spatulate thoracic notosetae and neuropodial thoracic SABELLINAE. Radioles in semi-circles; external
companion setae absent. Both thoracic and abdominal stylodes and webbing absent. Very large, compound
uncini avicular, with short, strongly bent handles. eyes present subdistally on a few dorsal radioli, other-
Fabricia Blainville 1828, Amphicora sabella Ehren- wise absent. Collar two- or four-lobed. Spatulate tho-
berg 1837; 18 species. racic notosetae present. Abdominal uncini avicular.
FABRICINAE. Three pairs of radioli and branchial Monroika Hartman 1951 b, Manayunkia africana Monro
hearts present. Collar often very low dorsally and 1939b; only species.
united ventrally. Abdomen with long-handled uncini. FABRICINAE. Six pairs of radioles, in part united
Three abdominal setigers. by a web. Collar ventrally high, dorsally incised.
Fabriciola Friedrich 1939, Manayunkia pacifica An- Vascularized filaments absent. Two abdominal setigers;
nenkova 1934; 6 species. abdominal uncini long-handled.
FABRICINAE. Three pairs of radioles and branchial Myxicola Koch in Renier 1847, Terebella infundibulum
hearts present. Collar distinct dorsally, united ven- Renier 1804; 5 species.
trally. Abdominal uncini short-handled. Three ab- MYXICOLINAE. Radioles in semi-circles, strongly
dominal setigers. webbed, external stylodes and eyes absent. Thoracic
Fabrisabella Hartman 1969, F. vasculosa Hartman uncini minute and long-handled. Abdominal uncini
1 969; 2 species. avicular, in nearly complete cinctures around the
FABRICINAE. Radioles in semi-circles; webbed, posterior part of the body. Tube mucoid.
eyes and external stylodes absent. Collar high, but Oriopsis Caullery and Mesnil 1896, Fabricia armandi
deeply and widely separated dorsally. Spatulate thoracic Claparede 1864; 22 species.
notosetae present; thoracic neuropodial companion FABRICINAE. Three to five pairs of radioles;
setae absent. Thoracic uncini long-handled and gently radioles externally flanged, webbing and eyes absent.
curved; abdominal ones strongly bent. Numerous pairs Collar divided dorsally and fused ventrally. Abdomen
of radioles present; abdomen with numerous setigers. with four or more setigers; abdominal uncini with short,
Hypsicomus Grube 1870b, Sabella phaeotaenia quadrangular handles and teeth in several rows along
Schmarda 1861; 12 species. one margin.
SABELLINAE. Radioles in semi-circles; webbing, Panousea Rullier and Amoureux 1970, P. africana
and external stylodes absent; eyes present. Collar well Rullier and Amoureux 1970; only species.
developed. Spatulate thoracic notosetae present; first SABELLINAE. Radioles in semicircles. Webbing,
notosetae in a long, straight or gently curved row. eyes and external stylodes absent. Collar four-lobed.
Abdominal uncini avicular. Thoracic uncini long-shafted, neuropodial companion
Jasmineira Langerhans 1880, J. caudata Langerhans setae present.
1880; 13 species.
Potamethus Chamberlin 1919c, Potamilla malmgreni
FABRICINAE. Eight or more pairs of radioli; web- Hansen 1878; 10 species.
bing and eyes absent. Collar well developed. Abdominal
SABELLINAE. Radioles in semi-circles; eyes, ex-
uncini avicular; several abdominal setigers present.
ternal stylodes and webbing absent. Collar very low,
Laonome Malmgren 1866, L. kroeyeri Malmgren 1866; except ventrally where it is produced into a pair of
4 species. triangular lobes. Thoracic uncini with long, nearly
SABELLINAE. Radioles in semi-circles; external straight handles; abdominal uncini short-stemmed.
stylodes and webbing absent; eyes usually present. Companion setae present.
Collar bilobed. Spatulate thoracic notosetae present;
Potamilla Malmgren 1866, Sabella neglecta Sars
thoracic neuropodial companion setae absent. Both
1 850; 30 species.
thoracic and abdominal uncini with short, flattened
base and avicular head. SABELLINAE. Radioles in semi-circles; external
stylodes and webbing absent; eyes present. Collar
Manayunkia Leidy 1859, M. speciosa Leidy 1859; 10 two- or four-lobed. Spatulate thoracic notosetae pres-
species. ent. Thoracic uncini short-handled.
FABRICINAE. Two pairs of radioli, ventrally in ad-
Potaspina Hartman 1969, P. pacifica Hartman 1969;
dition a pair of palplike, vascularized smooth filaments.
only species.
Webbing absent. Collar well developed; abdomen with
three setigers; abdominal uncini long-handled. SABELLINAE. Radioles in semi-circles; eyes, ex-
ternal stylodes and webbing absent. Collar low dorsally,
Megalomma Johansson 1926, Amphitrite vesiculosa high and bifid ventrally. Anterior thoracic neuropodia
Montagu 1815; only species. with avicular uncini; last thoracic neuropodia with

thick, acicular spines. Spatulate setae and companion this genus from all other FABRICIIN genera is incor-
setae present. Abdominal uncini small, acicular. rect; it does not have the webbed tentacular crown
claimed by Hartman. Members of this genus will key
Pseudobranchiomma Jones 1962, P. emersoni Jones out with Manayunkia above. It differs from the latter
1962; only species.
in that it has short limbate setae in both the first setigers;
SABELLINAE. Radioles in semi-circles; eyes ab-
Manayunkia has such setae only in the first setiger
sent, webbing absent; external stylodes present, but
and they are much longer. Furthermore, Monroika
small. Thoracic uncini short-handled, companion setae
has two rather than three abdominal setigers. Jones
absent. Four thoracic setigers.
(1974a) also described a new genus, Brandtika (geno-
Pseudofabricia Cantone 1972, P. aberrans Cantone type: B. asiatica), which will key out in the same
1972; only species. complex. It differs from other genera in this complex
FABRICINAE. Eight radioles on a greatly pro- in that it has pilose setae in the last three thoracic
longed, distally bifid anterior end and an extended setigers. This genus could not be included in the key,
rounded, lower lip. Abdominal uncini with short base since it was based on dried material, and no information
and several rows of teeth. Three abdominal setigers. was available about the structure of the tentacular
crown.
Sabella Linnaeus 1767, S. penicillus Linnaeus 1767;
35 species.
Invalid Genera
SABELLINAE. Radioles in semi-circles; eyes pres-
ent, external stylodes and webbing absent. Collar Amphicora Ehrenberg 1837, see Fabricia
four-lobed. Spatulate thoracic notosetae absent. Amphicorina Quatrefages 1866, see Oriopsis
Anamobaea Kroyer 1856, see Hypsicomus
Sabellastarte Savigny 1818, Eurato sanctijosephi
Arippasa Johnston 1865, see Myxicola
Gravier 1906; 8 species.
Aspeira Bush 1904a, see Potamilla
SABELLINAE. Radioles spiralled; eyes present,
Branchiomma Claparede 1870a, see Megalomma
webbing and external stylodes absent. Collar well
Chaponella Rullier 1972, see Euchone
developed, widely separated dorsally. Spatulate and
Dasychone Sars 1862, see Branchiomma
companion setae absent.
Dasychonopsis Bush 1904a, see Branchiomma
Schizobranchia Bush 1904a, S. insignis Bush 1904a; Distylia Quatrefages 1866, see Bispira
2 species. Dybowscella Nusbaum 1901, see Manayunkia
SABELLINAE. Radioles in semi-circles; dichoto- Eriographis Grube 1850, see Myxicola
mously divided. Eyes present; external stylodes and Eurato Saint-Joseph 1894, see Hypsicomus
webbing absent. Collar four-lobed. Thoracic uncini Fabriciella Zenkevitch 1935, see Fabriciola
long-handled, but bent; companion setae present. Garjaiowella Dybowskii 1929, see Manayunkia
Gorbunovia Annenkova 1952, see Potamethus
Spirographis Viviani 1805, S. spallanzani Viviani
Gymnosoma Quatrefages 1866, see Myxicola
1805, 4 species.
Haplobranchus Bourne 1883, see Manayunkia
SABELLINAE. Radioles spiralled, with one half
Hypsicomatides Augener 1922, see Hypsicomus
very much larger than the other, only one part spiralled.
Hypsicomatopsis Augener 1922, see Hypsicomus
Stylodes and webbing absent; eyes present. Collar
Laonomedes Chamberlin 1919c, see Potamilla
four-lobed. All uncini avicular.
Leiobranchus Quatrefages 1850, see Myxicola
Leptochone Claparede 18706, see Myxicola
Taxonomic Notes
Megachone Johnson 1901, see Chone
The sub-families as accepted here are based on the Metachone Bush 1904a, see Chone
structure of the neuropodial uncini of the thorax, in Metalaonome Bush 1904a, see Bispira
that members of the FABRICINAE have gently curved, Notaulax Tauber 1879, see Hypsicomus
long-handled uncini, and members of SABELLINAE Oria Quatrefages 1866, see Oriopsis
have strongly bent, avicular uncini. However, all Oriades Chamberlin 1919c, see Oriopsis
sabellids with thoracic neuropodial companion setae Oridia Rioja 1917, see Oriopsis
have been included in the SABELLINAE, irrespective Othonia Johnston 1835, questionably Fabricia
of the structure of the uncini. Parachonia Kinberg 1867b, see Chone
The genus Pseudopotamilla Bush is considered here Parasabella Bush 1904a, see Demonax
a junior synonym of Potamilla and the genus Tri- Potamis Ehlers 1887, see Potamethus
chosobranchella Dybowski 1929, is referred to Protulides Webster 1884, see Hypsicomus
Manayunkia. Pseudopotamilla Bush 1904a, see Potamilla
Jones (1974a) revised the original material of Mon- Sabellina Dujardin 1839a, indeterminable
roika and pointed out that the character used to separate Sabina Williams 1851, indeterminable

Sitophaga Gistel 1848, see Fabricia The family is known for a single genus and species,
Trichosobranchella Dybowski 1929, see Manayunkia Sabellonga disjuncta Hartman 1969 from northern
Baja California. The specimen resembles a sabellid
that has lost its tentacular crown. However, there is
FAMILY CAOBANGIIDAE JONES 1974b no trace of the loss of a tentacular crown, or that such
Small, short-bodied sabelliform polychaetes with has ever been present. The presence of the giant falcate
three pairs of radioli in a tentacular crown. Digestive spines in far posterior setigers also is characteristic.
tract U-shaped with the anus opening dorsally and far Otherwise, the setal equipment is largely what one
anteriorly on the body. First setiger with neurosetal would expect in a member of the SABELLINAE.
palmate hooks, remainder of the thoracic setigers with-
out hooks; two kinds of avicular hooks present in a FAMILY SERPULIDAE JOHNSTON 1865
posterior region. Body separated into two regions; a thorax with a
This family was recently proposed by Jones (1974b) thoracic membrane (absent in rare instances) and dorsal
for a series of small polychaetes that live in close capillary or limbate setae and an abdomen with ventral
association with molluscs in freshwater in Southeast setae and dorsal uncini. One radiole often transformed
Asia. Most of the forms burrow in the shell of the into an operculum. Tube calcareous.
host. The family consists of a single genus, Caobangia The serpulids and the closely allied spirorbids repre-
Giard 1893, with type-species C. billeti Giard 1893 sent a sub-specialty of their own within the polychaetes.
and a total of seven known species. Up to this time, The family has been reviewed on a couple of occasions;
the genus has been considered among sabellid enig- Saint-Joseph (1894) published an extensive review and
maticae, but the demonstration of a series of forms Southward (1963) gave a key to all known genera.
spread over a larger geographical region, made the The generic sub-divisions within the family are never-
recognition of a new family necessary. theless debatable. The older taxonomic groupings
placed emphasis on the structure of the operculum
and on the overall body-construction (development of
FAMILY SABELLONGIDAE HARTMAN 1969
the thoracic membrane and collar, presence or absence
Body cylindrical with few thoracic and many ab- of setae in specified segments, etc.). More recently,
dominal segments. Peristomium forms a bevelled some authors (especially Zibrowius) have placed more
collar around the prostomium (tentacular crown absent). emphasis on the detailed structure of the uncini and
Setae include long-handled uncini with companion setae. The key below represents a compromise, and
setae and giant falcate spines. may, as such, be difficult to use.
Key to Genera
Ia. Operculum absent 2
lb. Operculum present 7
2a (Ia). Abdominal setae trumpet-shaped Pseudoserpula
2b (Ia). Abdominal setae marginally dentate and geniculate, limbate or capillary 3
3a (2b). Slender capillary or limbate collar setae present 4
3b (2b). Collar setae modified 6
4a (3a). Thoracic membrane absent Salmacinopsis
4b (3a). Thoracic membrane present 5
5a (4b). Thoracic membrane reaches setiger 5 Subprotula
5b (4b). Thoracic membrane reaches setiger 7 Salmacina
6a (3b). Abdominal setae geniculate Salmacina
6b (3b). Abdominal setae slender, nearly straight capillaries Protis
7a (Ib). Operculum carried on a branchial radiole 8
7b (ib). Operculum carried on a modified stalk 12
8a (7a). Five thoracic setigers present 9
8b (7a). Six or more thoracic setigers present 10
9a (8a). Collar setae slender and limbate; long asetose region present between thorax and abdomen
Josephella
9b (Sa). Collar setae basally dentate; thorax and abdomen not separated by a long asetose region
Dipomatus
l0a (8b). Collar setae simple and tapered limbate Appmatus

l0b (8b). Collar setae with a few coarse teeth basally and a limbate denticulate blade distally 11
I la (l0b). Operculum a shallowly depressed cone on a spherical swelling of the opercular stalk ... Filogranula
I lb (l0b). Operculum a depressed cone; stalk without the subdistal swelling Filograna
12a (7b). Opercular stalk flattened and ribbonlike Metavermilia
t2b (7b). Opercular stalk oval or circular in cross-section (sometimes with wings or spines) 13
13a (12b). Opercular stalk with wings or spines 14
13b (12b). Opercular stalk without wings or spines 26
14a (13a). Opercular stalk with spines 15
14b (13a). Opercular stalk with wings (sometimes produced into points distally) 17
15a (l4a). Opercular stalk with spines on one side only; six thoracic setigers present Spirodiscus
15b (14a). Opercular stalk with four spines in a cross; seven thoracic setigers present 16
16a (15b). Thoracic neuropodia widely separated anteriorly and approaching posteriorly; collar setae all of one
kind, with basal pilose boss and distal spines Spirobranchus
16b (15b). Thoracic neuropodia equidistant in all setigers; collar setae of two kinds; either capillary or with a
smooth double-boss basally and smooth tapering tips Crucigera
17a (13a). Collar setae absent 18
17b (13a). Collar setae present 19
18a (17a). Operculum with two projections Olga
l8b (17a). Operculum without projections Pomatoleios
19a (17b). Collar setae limbate or capillary 20
19b (17b). Collar setae with two separate limbations (bayonet-type), with a basal boss or setose 23
20a (19a). Anterior abdominal setae stout, acute spines Paumotella
20b (19a). Anterior abdominal setae trumpet-shaped or geniculate 21
21a (20b). Opercular cap black and chitinous Crosslandiella
21b (20b). Opercular cap calcareous 22
22a (21b). Opercular cap flat, with or without spines Pomatoceros
22b (21b). Operculum distally excavated, bordered by two eccentrically placed thorns connected by a low
ridge Pseudopomatoceros
23a (19b). At least anterior abdominal setae trumpet-shaped 24
23b (19b). Anterior abdominal setae geniculate 25
24a (23a). Operculum conical, collar setae basally pilose Conopomatus
24b (23a). Operculum with slanting distal plate, collar setae bayonet-shaped Temporaria
25a (23b). Uncini with anterior peg pointed Omphalopomopsis
25b (23b). Uncini with anterior peg gouge-shaped (hollowed out from beneath) Pomatostegus
26a (13b). Collar setae absent 27
26b (13b). Collar setae present 33
27a (26a). Abdominal setae absent; abdominal uncini present Rhodopsis
27b (26a). Abdominal setae present; uncini usually present 28
28a (27b). At least some abdominal setae geniculate 29
28b (27b). All abdominal setae slender capillaries 30
29a (28a). Uncini with numerous teeth Placostegus
29b (28a). Uncini with seven to nine teeth Marifugia
30a (28b). Abdominal uncini absent Bonhourella
30b (28b). Abdominal uncini present 31
31a (30b). Tube free, tusk-shaped and smooth Ditrupa
31b (30b). Tube at least partly attached 32
32a (31b). Opercular stalk calcified; operculum funnel-shaped Sclerostyla (in part)
32b (31b). Opercular stalk not calcified; operculum spherical Dasynema
33a (26b). Opercular spines movable . . . . . . . . . . . . . . . . . . . . . . . . . . .......................... Galeolaria
33b (26b). Opercular spines, if present, immovable 34
34a (33b). Abdominal setae slender capillaries Schizocraspedon
34b (33b). Abdominal setae trumpet-shaped or geniculate 35
35a (34b). Abdominal setae trumpet-shaped 36
35b (34b). Abdominal setae geniculate 39
36a (35a). Operculum a simple funnel 37

/976 THE POLYCHAETE WORMS /43
36b (35a). Operculum with a basal funnel and in addition various spines or hoods forming a distal part .... 38
37a (36a). Collar setae basally minutely hirsute Paraserpula
37b (36a). Collar setae basally dentate Serpula
38a (36b). Operculum with a large, glandular hood-shaped distal part Olgaharmania
38b (36b). Operculum with series of distal spines and, sometimes, smaller hoods Hydroides
39a (35b). Sicle setae (Apomatus-setae) absent 40
39b (35b). Sicle setae present 49
40a (39a). Collar setae with few coarse teeth in one or two marginal rows 41
40b (39a). Collar setae at least in part limbate 45
41a (40a). Operculum distally ornamented 42
41b (40a). Operculum distally smooth, rounded or excavate 43
42a (41 a). Operculum with concentric series of teeth, nearly radial in structure Neopomatus
42b (41 a). Operculum with a single series of long, strong teeth, making it bilaterally symmetrical . Mercierella
43a (416). Collar setae with teeth in two rows Sphaeropomatus
43b (41b). Collar setae with teeth in a single row 44
44a (43b). Operculum distally rounded Ficopomatus
44b (43b). Operculum distally excavate Merciere/lopsis
45a (40b). At least some collar setae with a basal dentate or hirsute region 46
45b (40b). All collar setae limbate 48
46a (45a). Simple limbate collar setae present, in addition to some with a basal group of spines Omphalopoma
46b (45a). All collar setae with basal denticulated or hirsute region 47
47a (46b). Six thoracic setigers present Hyalopomatus
47b (46b). Seven thoracic setigers present Pseudochitinopoma
48a (45b). Opercular stalk calcified; abdominal setae with only a short geniculate tip Sclerostyla (in part)
48b (45b). Opercular stalk not calcified; abdominal setae with about one-half of the exposed setae in the
geniculate tip Neovermilia
49a (39b). Opercular stalk annulated Calcareopomatus
49b (39b). Opercular stalk not annulated 50
50a (49b). Collar setae limbate 51
50b (49b). Collar setae with a basal boss in addition to the limbate distal portion 52
51a (50a). Both thoracic and abdominal uncini with all teeth in a single row; anterior peg entire . . Vermiliopsis
51b (50a). Thoracic uncini with teeth in a single row, anterior peg furcate; abdominal uncini with teeth in
several rows Pseudovermilia
52a (51 a). Both thoracic and abdominal uncini with teeth in several rows Filogranula (in part)
52b (51 a). Thoracic uncini with teeth in a single row 53
53a (52b). Operculum distally rounded Chitinopomoides
53b (52b). Operculum distally excavate 54
54a (53b). Opercular stalk with three distal bulbs Janita
54b (53b). Opercular stalk without swellings Chitinopoma
Generic Definitions absent. Thoracic uncini with anterior peg gouge-

Apornatus Philippi 1844, A. ampuiliferus Philippi shaped.
1844; 8 species. Calcareopomatus Straughan 1967a, C. dewae Straughan
SERPULINAE. Seven thoracic setigers; globular 1 976a; only species.
operculum on a radiole. Collar setae simple limbates; SERPULINAE. Seven thoracic setigers; operculum
sicle setae present. Uncini with teeth in several rows with a globular base and a flat, calcareous plate; stalk
in both thorax and abdomen; anterior peg very long. annulated. Collar setae small and limbate; side setae
Bonhourella Gravier 1905b, B. insignis Gravier 1905b; present. Thoracic uncini with anterior peg bifid.
only species. Chitinopoma Levinsen 1884, Hydroides norvegica
SERPULINAE. Six thoracic setigers; operculum flat groenlandica Morch 1863; 2 species.
or lightly convex, opercular stalk distinct. Collar setae SERPULINAE. Seven thoracic setigers; operculum
absent; abdominal setae capillary; abdominal uncini conical without wings or spines on the stalk. Collar
I

setae of fin and blade construction; sicle setae present; FICOPOMATINAE. Seven thoracic setigers; oper-
abdominal setae geniculate with triangular blade. culum pear-shaped, soft or chitinous; stalk smooth.
Thoracic uncini with teeth in a single row; abdominal Some collar setae with dentate boss, others serrated
ones with teeth in several rows. limbates. Sicle setae absent. Abdominal setae genicu-
late with a dentate, slender tip. Uncini with few teeth
Chitinopomoides Benham 1927, C. wilsoni Benham
in a single row.
1927; only species.
SERPULINAE. Seven thoracic setigers; operculum Filograna Oken 1815, F. implexa Berkeley 1828; only
rounded, stalk without spines or wings. Collar setae species.
with large boss below a limbate zone; sicle setae pres- FILOGRANINAE. Six to 12 thoracic setigers;
ent; abdominal setae geniculate with long triangular rounded operculum on one of the radioles. Collar setae
blade. Thoracic uncini with teeth in a single row; notched with limbate expansion at the base. Sicle setae
abdominal ones with teeth in several rows. and limbate setae present in thorax; abdomen with
geniculate setae. Uncini with numerous teeth in several
Conopomatus Pillari 1960, C. acuiconus Pillai 1960; rows.
2 species.
SERPULINAE. Seven thoracic setigers; operculum Filogranula Langerhans 1884, F. gracilis Langerhans
conical, stalk winged. Collar setae present, pilose at 1884; 3 species.
base; side setae present; other thoracic setae limbate; FILOGRANINAE. Seven thoracic setigers; opercu-
abdominal setae trumpet-shaped. lum a small cone on a small, spherical base; stalk
either frondose or smooth; only four radioles present.
Crosslandiella Monro 1933, C. multispinosa Monro Collar setae present; side setae present; uncini with
1933; 2 species. numerous teeth in several rows.
SERPULINAE. Seven thoracic setigers; operculum
with black, chitinous plate, surmounted by a spinose col- Galeolaria Savigny 1818, G. caespitosa Savigny 1818;
umn; stalk winged. Collar setae lancet-shaped and taper- 5 species.
ing; side setae present. Abdominal setae geniculate. SERPULINAE. Seven thoracic setigers; operculum
with calcareous plate surmounted by movable spines;
Crucigera Benedict 1887, C. websteri Benedict 1887; opercular stalk winged. Collar setae present and very
5 species. short and slender.
Hyalopomatus Marenzeller 1878, H. claparedii Maren-
with simple funnel, distal end of stalk with four large
zeller 1878; 4 species.
spines forming a cross. Collar setae limbate; side setae
SERPULINAE. Six thoracic setigers; operculum
absent; abdominal setae trumpet-shaped. Uncini with
bladder-shaped; stalk smooth. Collar setae of fin and
few teeth; anterior peg very large.
blade construction; side setae absent. Abdominal setae
Dasynema Saint-Joseph 1894, Serpula chrysogyrus with only the distalmost tip geniculate. Uncini with
Grube 1878; only species. teeth in several rows; anterior peg furcate.
Hydroides Gunnerus 1768, H. norvegica Gunnerus
spherical, opercular stalk smooth. Collar setae and
1768; 85 species.
side setae absent; all thoracic setae limbate, abdominal
ones capillaries.
basally with a marginally dentate funnel, distally with
Dipoinatus Ehlers 1913, D. serpulides Ehlers 1913; a crown of spines or smaller hoods, or a second, distal
only species. funnel present; stalk without spines or wings. Collar
SERPULINAE. Five thoracic setigers; two operculae setae limbate; side setae absent; abdominal setae
with branchial filaments on peduncle; each distally trumpet-shaped.
funnel-shaped and marginally dentate. Collar-setae
Janita Saint-Joseph 1894, Serpula fimbriata delle
basally dentate. Chiaje 1828; only species.
Ditrupa Berkeley 1835, Dentalium arietinum O.F. SERPULINAE. Seven thoracic setigers; operculum
Muller 1776; 8 species. infundibular with three large distal bulbs on the stalk;
SERPULINAE. Tube free, tusk-shaped; operculum opercular plate chitinous with a single large spine.
an inverted cone with a chitinous plate. Collar setae Collar setae basally dentate; side setae present; ab-
absent; thoracic setae limbate and capillaries; abdominal dominal setae geniculate.
setae capillaries. Uncini with numerous teeth in several
Josephella Caullery and Mesnil 1896, J. marenzelleri
rows. Caullery and Mesnil 1896; 2 species.
Ficopomatus Southern 1921, F. macrodon Southern SERPULINAE. Five thoracic setigers; operculum
1921; 2 species. at end of normal radiole, rounded. Collar setae slender

limbate; uncini with double rows of teeth; anterior SERPULINAE. Six thoracic setigers; operculum
peg deeply bifid. with a central tooth and paired lateral horns, stalk
winged. Collar setae absent, side setae absent; tho-
Marifugia Absalon and Hrabe 1930, M. cavatica
racic setae limbate; uncini with about twenty teeth;
Absalon and Hrabe 1930; only species.
anterior peg gouge-shaped.
SERPULINAE. Six thoracic setigers; operculum
conical and distally smooth; stalk without spines or Olgaharmania Rioja 1941b, Hydroides glandiferum
wings. Collar setae and side setae absent; thoracic Rioja 1941a; only species.
setae slender and straight; abdominal setae geniculate. SERPULINAE. Seven thoracic setigers; operculum
Uncini with teeth in a single row, maximally nine with basal corona of spines surmounted by very large,
teeth present in thoracic uncini. glandular chitinous hood equipped on one side with
Membranopsis Bush 1910,M. inconspicua Bush 1910; four large spines. Collar setae basally with paired
only species. bosses; side setae absent. Uncini with numerous teeth;
SERPULINAE. Nine thoracic setigers; operculum anterior peg very large.
unknown. Thoracic membrane fused over the back at Omphalopoma Morch 1863, 0. umbilicata Morch
ninth setiger; collar setae limbate. Uncini similar to 1863; 2 species.
those in Protula. SERPULINAE. Six or seven thoracic setigers; oper-
Mercierella Fauvel 1923b, M. enigmatica Fauvel culum cup-shaped, sometimes ornamented; stalk smooth.
1923b; only species. Collar setae of two kinds, simple limbates and fin and
FICOPOMATINAE. Seven thoracic setigers; opercu- blade setae. Side setae present. Thoracic uncini with
lum distally oblique with chitinous plate covered by teeth in a single row; abdominal ones with teeth in
series of curved spines. Collar setae marginally dentate; several rows; anterior peg furcate.
side setae absent; uncini with teeth in one row in thorax,
Omphalopomopsis Saint-Joseph 1894, Omphalopoma
partially in two rows in abdomen.
langerhansii Marenzeller 1884; 2 species.
Mercierellopsis Rioja 1945, M. prietoi Rioja 1945; SERPULINAE. Seven thoracic setigers. Operculum
only species. distally smooth; opercular stalk with wings. Collar
FICOPOMATINAE. Seven thoracic setigers; opercu- setae of two kinds, slender capillaries and limbate
lum smooth, distally cup-shaped; stalk smooth. Collar with a pilose subdistal boss. Sicle setae present; ab-
setae dentate; side setae absent. Uncini with few teeth dominal setae geniculate with a triangular blade.
(7-12), anterior peg undercut. Thoracic uncini with teeth in a single row; anterior
peg entire.
Metavermilia Bush 1904a, Vermilia multicristata
Philippi 1844; 4 species. Paraserpula Southward 1963, P. planorbis Southward
SERPULINAE. Seven thoracic setigers; operculum 1963; only species.
bladder-shaped; stalk flattened, ribbon-shaped. Collar SERPULINAE. Seven thoracic setigers; operculum
setae limbate; side setae present; abdominal setae funnel-shaped, marginally dentate; stalk smooth. Collar
geniculate or capillaries. Thoracic uncini with teeth setae bayonet-shaped and finely pilose; side setae
in a single row; abdominal ones with teeth in one or absent; abdominal setae trumpet-shaped. Thoracic
several rows. uncini with teeth in a single row; abdominal ones with
teeth in several rows; anterior peg large.
Neopomatus Pillai 1960, N. uschakovi Pillai 1960;
2 species. Paumotella Chamberlin 1919c, P. takemoana Cham-
FICOPOMATINAE. Seven thoracic setigers; opercu- berlin 1919c; only species.
lum with series of concentric rows of teeth, nearly SERPULINAE. Seven thoracic setigers present;
radial in structure; stalk smooth. Collar setae dentate; operculum conical and distally flattened, margin entire,
side setae absent. Uncini with few teeth. stalk with wings. Collar setae limbate; side setae ab-
Neovermilia Day 1961, N. capensis Day 1961; 2 sent; anterior abdominal setae stout acute spines; pos-
species. terior ones fine capillaries.
Placostegus Philippi 1844, Serpula tridentata Fabricius
spherical or slightly cup-shaped; stalk smooth. Collar
1 780; 18 species.
setae limbate; side setae absent. Abdominal setae
geniculate, finely dentate capillaries. Thoracic uncini
funnel-shaped with chitinous distal plate; stalk smooth.
with few teeth in a single row.
Collar setae absent; abdominal setae geniculate with
Olga Jones 1962, 0. elegantissima Jones 1962; only widely triangular dentate distal plates. Uncini with
species. numerous teeth.

Pomatoceros Philippi 1844, Serpula triquetra Linnaeus Pseudoserpula Straughan 1967b, P. rugosa Straughan
1767; 13 species. 1967b; 2 species.
SERPULINAE. Seven thoracic setigers; operculum SERPULINAE. Seven thoracic setigers; operculum
a flat calcareous plate, with or without spines; stalk absent. Collar setae limbate; side setae absent; ab-
with broad wings. Collar setae small and limbate; dominal setae trumpet-shaped. Uncini with few teeth;
thoracic setae limbate; abdominal setae trumpet-shaped. anterior peg large, blunt.
Sicle setae absent. Uncini with numerous teeth.
Pseudovermilia Bush 1907b, Spirobranchus occidentalis
Pomatoceios Pixell 19136, P. crosslandi Pixell 19136; McIntosh 1885; 5 species.
3 species. SERPULINAE. Seven thoracic setigers; operculum
SERPULINAE. Seven thoracic setigers; operculum distally ornamented; stalk smooth. Collar setae limbate,
distally flat; stalk winged. Collar setae and side setae side setae present; abdominal setae geniculate with
absent; abdominal setae trumpet-shaped with one long triangular blades. Thoracic uncini with teeth in a single
spine at one side. Uncini with about ten teeth in both row; anterior peg furcate; abdominal ones with teeth
thorax and abdomen. in several rows.
Pomatostegus Schmarda 1861, P. macrosoma Schmarda Rhodopsis Bush 1904a, R. pusillus Bush 1904a; only
1861; 8 species. species.
SERPULINAE. Seven thoracic setigers; operculum SERPULINAE. Seven thoracic setigers; operculum
with chitinous plate surmounted by a column with covered distally with chitinous plate with spines ar-
several discs; stalk winged. Collar setae with subdistal ranged in a rosette; shaft smooth. Collar setae absent;
pilose boss; side setae present; abdominal setae genicu- abdominal setae absent. Uncini with teeth in a single
late. Uncini with teeth in a single row; anterior peg row.
simple.
Salmacina Claparede 1870a, S. incrustans Claparede
Protis Ehlers 1887, P. simplex Ehlers 1887; 3 species. 1870a; 8 species.
SERPULINAE. Seven thoracic setigers; operculum FILOGRANINAE. Five to nine thoracic setigers;
absent. Collar setae of fin and blade construction with operculum absent. Collar setae notched with limbate
basal dentate fin well separated from the blade. Sicle subdistal and distal parts; side setae present. Abdominal
setae present; abdominal setae nearly capillary. Tho- setae geniculate. Uncini with teeth in several rows.
racic uncini with teeth in a single row; abdominal ones
Salmacinopsis Bush 1910, S. setosa Bush 1910; only
with teeth in several rows.
species.
Protula Risso 1826, Serpula tubularia Montagu 1803 FILOGRANINAE. Nine thoracic setigers; operculum
in McIntosh 1923; 23 species. absent. Collar setae capillary; other thoracic setae
SERPULINAE. Seven thoracic setigers; operculum capillary and limbate with a few serrate setae in pos-
absent. Collar setae simple limbates; side setae pres- terior thoracic setigers; abdominal setae geniculate
ent. Uncini with teeth in several rows; anterior peg and serrated. Uncini with numerous very fine teeth;
very long. anterior peg large and blunt. Thoracic membrane
absent.
Pseudochitinopoma Zibrowius 1969, Hyalopomatopsis
occidentalis Bush 1904a; only species. Schizocraspedon Bush 1904a, Serpula furcifera Grube
SERPULINAE. Seven thoracic setigers; operculum 1 878; only species.
bladder-shaped; stalk without wings. Collar setae with SERPULINAE. Seven thoracic setigers; operculum
fin and blade construction and little separation between on a modified radiole; distally with a chitinous cover
the basal fin and the distal blade. Sicle setae absent; consisting of two concentric funnels, one inside the
abdominal setae geniculate with narrow blade. Tho- other; both marginally split with long, slender pro-
racic uncini with teeth in a single row; anterior peg cesses. Collar setae present. Abdominal setae capillary;
furcate; abdominal uncini with teeth in several rows. superior thoracic setae geniculate with conspicuous
spines at the base.
Pseudopomatoceros Holly 1936, Pomatoceropsis roxasi
Holly 1935; only species. Sclerosryla M6rch 1863, S. ctenactis Morch 1863;
SERPULINAE. Seven thoracic setigers; operculum 2 species.
distally cup-shaped with two eccentrically placed thorns SERPULINAE. Seven thoracic setigers; operculum
connected by a low ridge; stalk winged. Collar setae funnel-shaped; opercular stalk smooth, calcified, collar
fine capillaries; side setae absent, thoracic setae lim- setae present or absent, if present finely limbate; side
bate, abdominal setae with flat crown and a long spike. setae absent; abdominal setae nearly capillary, with
Uncini with numerous teeth; anterior peg furcate. very short geniculate tips. Thoracic uncini with teeth

in a single row; abdominal ones with teeth in several Taxonomic Notes

rows.
The genus Membranopsis Bush 1910, included in
Serpula Linnaeus 1767, S. vermicularis Linnaeus the generic definitions, has been omitted from the key,
1 767; 17 species. since the structure of the operculum remains unknown.
SERPULINAE. Seven thoracic setigers; operculum The genus Hydroides has been defined here to in-
funnel-shaped with crenulated margin; opercular stalk clude Eupomatus Philippi 1844, as is customarily done
smooth. Collar setae either limbate or with a dentate by most specialists today. It may be of some value to
boss; side setae absent. Abdominal setae trumpet- recognize the difference between Hydroides SeNsu
shaped. Uncini with few teeth; anterior peg large. STRICTO and Eupomatus at the sub-generic level.
Sphaeropomatus Treadwell 1934, S. miamiensis

Treadwell 1934; only species. Invalid Genera
FICOPOMATINAE. Seven thoracic setigers; opercu-
l um soft and vesicular; stalk smooth. Collar setae Apomatopsis Saint-Joseph 1894, see Apomatus
spinose with teeth in two rows; side setae absent; Cystopomatus Gravier 191 Ia, see Hyalopomatus
other setae slender capillaries. Eucarphus Morch 1863, see Hydroides
Eupomatus Philippi 1844, see Hydroides
Spirobranchus Blainville 1818, Serpula gigantea Pallas Filigrana Morch 1863, see Filograna
1 766; 30 species. Filopora Fleming 1825, see Filograna
SERPULINAE. Seven thoracic setigers; operculum Glossopsis Bush 1904a, see Hydroidess
with calcareous distal plate, usually ornamented; oper- Helena Castelnau 1842, see Serpula
cular stalk winged. Collar setae bayonet-shaped and Hyalopomatopsis Saint-Joseph 1894, see Hyalopo-
pilose; some forms have limbate collar setae; side matus
setae absent. Abdominal setae trumpet-shaped. Tho- Microserpula Dons 1931, see Chitinopoma
racic uncini with teeth in a single row; anterior peg Paravermilia Bush 1907b, see Vermiliopsis
furcate. Thoracic neuropodia are separated widely Piratesa Templeton 1835, indeterminable
anteriorly and approaching in late thoracic setigers. Placostegopsis Saint-Joseph 1894, see Placostegus
Spirodiscus Fauvel 1909, S. grimaldii Fauvel 1909; Podioceros Quatrefages 1866, see Poinatoceros
Polyphragma Quatrefages 1866, see Hydroides
only species.
Pomatoceropsi.s Gravier 1905b, see Spirobranchus
FILOGRANINAE. Six thoracic setigers; operculum
funnel-shaped on a small ampulla; stalk with dorsal Pomatoceropsis Holly 1 935, see Pseudopomatoceros
spines. Collar setae flattened and lancet-shaped. Uncini Proplacostegus Bush 1904a, see Placostegus
with numerous teeth. Psymobranchus Philippi 1844, see Protula
Siliquaria Savigny 1818, indeterminable
Subprotula Bush 1910, S. longiseta Bush 1910; 2 Spiramella Blainville in Quatrefages 1866, see Protula
species. Vermilia Savigny 1818, see Spirobranchus
SERPULINAE. Seven thoracic setigers; operculum Zopyrus Kinberg 1867b, see Serpula
absent. Collar setae capillaries. Uncini with about 15
teeth; anterior peg very large. Thoracic membrane to
fifth setiger only; collar tri-lobed. FAMILY SPIRORBIDAE PILLAI 1970
Temporaria Straughan 19676, Vermilia polytrema Body separated into two regions; a thorax with a
Philippi 1844; 2 species. thoracic membrane, dorsal setae and ventral uncini
SERPULINAE. Seven thoracic setigers; operculum and an abdomen with ventral setae and dorsal uncini.
with slanting calcareous plate; stalk winged. Collar One radiole transformed into an operculum; tube
setae bayonet-shaped with both regions limbate, sep- calcareous. Body asymmetrical and tube coiled in a
arated by a notch. Abdominal setae trumpet-shaped. spiral; thorax with setigerous rudiments of three, four
Thoracic uncini with at least ten teeth. or five segments.
The spirorbids are rather difficult to work with since
Vermiliopsis Saint-Joseph 1894, Serpula infundibulum identification requires precise and accurate work with
Linnaeus 1758; 35 species. compound microscopes; prior to that of course, the
SERPULINAE. Seven thoracic setigers; operculum specimens must be dissected out from their tubes as
rounded or flattened, stalk without wings. Collar and completely as possible. Recent reviews by Bailey-
sidle setae present. Thoracic uncini with teeth in a Brock, Knight-Jones and Vine has clarified the generic
single row; anterior peg entire; abdominal uncini with sub-division of the group considerably and is largely
teeth in a single row. followed here (references to articles can be found be-

low). Where the above-mentioned authors have yet The family is most frequently considered a sub-
to treat a genus, Pillai (1970) has been followed. family under the serpulids.
Key to Genera
Ia. With three or more pairs of thoracic tori 2

lb. With two pairs of thoracic tori 5
2a (1 a). With four or more pairs of thoracic tori or at least rudiments of a fourth pair present on the concave
side 3
2b (Ia). With three pairs of thoracic Lori 17
3a (2a). With at least four complete thoracic tori present 4
3b (2a). With remnants of a fourth thoracic torus present 21
4a (3a). Four pairs of thoracic tori present Anomalorbis
4b (3a). Approximately seven pairs of thoracic tori present Neomicrorbis
5a(lb). Sicle setae absent 6
5b (lb). Sicle setae present 10
6a (5a). Margins of collar fused to form dorsal tunnel Dexiospira
6b (5a). Margins of collar not fused 7
7a (6b). Dextrally coiled Pillaiospira
7b (6b). Sinistrally coiled 8
8a (7b). Collar setae limbate Leodora
8b (7b). Collar setae of fin and blade construction 9
9a (8b). Thoracic uncini with simple, blunt anterior peg Simplicaria
9b (8b). Thoracic uncini with broadly indented anterior peg Eulaeospira
10a (5b). Dextrally coiled 11
l0b (5b). Sinistrally coiled 12
I la (l0a). Collar setae of fin and blade construction Spirorbella
A
FIGURE 38. (A), Family SPIRORBIDAE, Laeospira sp., simplified after Day 1964, 15x; (B), Promlaeospira patagonica, tube,
after Day 1967, about 15x; (C), Family SERPULLDAE, Spirobranchus spinosuA, after Hartman, 1969, 5x; (D), Serpula ver-
micularis, tube, natural size.

I I b (l0a). Collar setae simply limbate Janua

12a (l0b). Collar setae simply limbate 13
12b (lob). Collar setae of fin and blade construction 15
13a (12a). Collar fused dorsally to form tunnel Romanchella
13b (12a). Collar not fused dorsally 14
14a (13b). Incubation of larvae in opercular chamber; abdominal setae with blades as those of the collar
setae Fauveldora
14b (13b). Incubation in the tube; abdominal setae very small with short blades Bushiella
15a (12b). Thoracic uncini with several rows of teeth Spirorbis
l5b (12b). Thoracic uncini with a single row of teeth 16
16a (15b). Two or more opercular plates on top of each other on the operculum Duplicaria
l6b (15b). A single opercular plate present Pileolaria
17a (2b). Sicle setae present 18
17b (2b). Sicle setae absent 20
18a (17a). Collar setae of fin and blade construction Paralaeospira
18b (17a). Collar setae simply limbate 19
19a (18b). Incubation of larvae in opercular chamber Protoleodora
l9b (18b). Larvae attached to stalk in faecal groove Metalaeospira
20a (17b). Tube tightly coiled along its whole length Circeis
20b (17b). Tube coiled only basally, most of the tube uncoiled Helicosiphon
21 a (3b). Three thoracic tori and four fascicles of setae on the convex side; four thoracic tori and five fascicles
of setae on the concave side Amplaria
21b (3b). Fourth setiger represented by a rudiment of the torus on the concave side 22
22a (21b). Sinistrally coiled 23
22b (21b). Dextrally coiled 24
23a (22a). Sicle setae present Protolaeospira
23b (22a). Sicle setae absent Capeospira
24a (22b). Collar setae simply limbate Prodexiospira
24b (22b). Collar setae of fin and blade construction 25
25a (24b). Thoracic uncini with furcate anterior peg Dextralia
25b (24b). Thoracic uncini with bluntly rounded anterior peg Paradexiospira
The key given above suggests that all named taxa generic level, as indicated in Table 3. 1 believe that
are at the same level. Vine (1972) and Knight-Jones this difference in approach is relatively unimportant
(1973) used several of the contained taxa at the sub- for the time being.
TABLE 3
Alphabetic list of genera and sub-genera of Spirorbidae as suggested by
Vine (1972) and Knight-Jones (1973).
Genus Amplaria Genus Paralaeospira

Anomalorbis Pileolaria
Capeospira subgenus Duplicaria
Circeis Pileolaria
Eulaeospira Protoleodora
Helicosiphon Simplicaria
JanUn Prodexiospira
subgenus Dexiospira Protolaeospira
Fauveldora subgenus Dextralia
Janua Protolaeospira
Leodora Romanchella
Pillaiospira subgenus Bushiella
Metalaeospira Romanchella
Neomicrorbis Spirorbis
Paradexiospira subgenus Spirorbella
Spirorbis

Generic Definitions anterior peg. Abdominal setae with vestigial brush-

top. Incubation in faecal groove on attachment stalk.
Amplaria Knight-Jones 1973, A. spiculosa Knight- Duplicaria Vine 1972, Spirorbis (Laeospira) koehleri
Jones 1973; only species. Caullery and Mesnil 1897; 3 species.
Sinistrally coiling; four thoracic tori and four fas- Sinistrally coiled; two pairs of thoracic tori present.
cicles of setae on the concave side; three thoracic tori Collar setae of fin and blade construction, not cross-
and five fascicles of setae on the convex side. Fin striated. Sicle setae present; thoracic uncini with several
and blade collar setae without cross-striations; side rows of teeth. Anterior abdominal tori divided into
setae in third and fourth thoracic fascicle. Thoracic two parts. Operculum with two or more plates stacked
uncini with blunt anterior peg; abdominal setae genicu- on top of each other; incubation in the operculum.
late, less than one fourth the size of the sicle setae.
Abdomen somewhat asymmetrical; incubation in the Eulaeospira Pillai 1970, Spirorbis (Laeospira) orien-
operculum. talis Pillai 1960; 2 species.
Sinistrally coiled; two pairs of thoracic tori. Collar
Anomalorbis Vine 1972, A. manuatus Vine 1972; setae of fin and blade construction, without cross-
only species. striations. Sicle setae absent. Uncini with teeth in
Probably dextrally coiling; four complete thoracic several rows; anterior peg bifid or multifid. Incubation
setigers and remnants of a fifth. Collar setae limbate; i n groove along posterior abdomen.
side setae absent. Pronounced abdominal asymmetry;
Fauveldora Knight-Jones 1972, Janua (Fauveldora)
incubation probably in tube.
kayi Knight-Jones 1972; only species.
Bushiella Knight-Jones 1973, Spirorbis evolutus Bush Sinistrally coiled; two pairs of thoracic tori present;
1 9046; 4 species. collar setae limbate; abdominal setae with blades as
Sinistrally coiling; two pairs of thoracic tori present; big as those in the collar setae; side setae in third
collar setae limbate, numerous. Sicle setae present in fascicle present. Uncini with teeth in several rows.
third fascicle. Thoracic uncini with blunt anterior peg. Incubation in the operculum.
Margin of collar unfused; abdomen not strongly asym-
Helicosiphon Gravier 1907, H. biscoeensis Gravier
metrical. Incubation in faecal groove with attachment
1907; only species.
stalk for the embryos.
Sinistrally coiled, or almost uncoiled; three thoracic
Capeospira Pillai 1970, Spirorbis (Paralaeospira) setigers on concave side. Collar setae limbate; side
adeonella Day 1963; only species. setae absent. Thoracic uncini with blunt anterior peg.
Sinistrally coiling; thorax with three complete and Abdominal tori distinctly larger on concave side. In-
a fourth rudimentary setiger; collar setae of fin and cubation in faecal groove attached by stalk.
blade construction; side setae absent. Incubation in
Janua Saint-Joseph 1894, Spirorbis pagenstecheri
the tube. Quatrefages 1865; 7 species.
Circeis Saint-Joseph 1894, Serpula spirillum Linnaeus Dextrally coiled; two pairs of thoracic tori present;
1 758; 3 species. collar setae limbate; side setae in third fascicle present.
Dextrally coiled; three thoracic setigers present; Abdominal setae with blades as big as those in the collar
collar setae limbate. Sicle setae absent. Incubation seate. Uncini with teeth in several rows. Incubation
in the tube. in the operculum.
Dexiospira Caullery and Mesnil 1897, Serpula cor- Leodora Saint-Joseph 1894, Spirorbis laevis Qua-
rugata Montagu 1803; 23 species. trefages 1865; 8 species.
Usually dextrally coiled; two pairs of thoracic tori Sinistrally coiled; two pairs of thoracic tort present;
present. Collar setae limbate; abdominal setae with collar setae limbate, not dentate; side setae absent.
blades as long as those in the collar seta; side setae Abdominal setae with blades as big as those in the
absent. Margins of collar fused to form dorsal tunnel; collar setae. Uncini with teeth in several rows. Incu-
incubation in operculum. bation in the operculum.
Dextralia Knight-Jones 1973, Spirorbis (Paradexio- Metalaeospira Pillai 1970, Spirorbis pixelli Harris
spira) falklandica Pixell 1913a; only species. 1969; 4 species.
Dextrally coiled; three rows of tori on the concave Sinistrally coiled; three thoracic tori on the concave
side and traces of a fourth on the convex side. Collar side. Collar setae with limbate blades; side setae
setae of fin and blade construction; side setae in present. Both thoracic and abdominal uncini more
third and fourth fascicle. Thoracic uncini with bilobed numerous on concave side. Thoracic uncini with blunt


anterior peg. Incubation in faecal groove; attachment collar setae limbate; side setae present. Incubation in
stalk absent. operculum.
Neomicrorbis Roverto 1904, Serpula crenatostriatus Romanchella Caullery and Mesnil 1897, R. perrieri
Goldfuss 1833; 3 species. Caullery and Mesnil 1897; 4 species.
Sinistrally or dextrally coiled; approximately seven Sinistrally coiled; two pairs of thoracic tort present;
thoracic setigers. Collar setae of fin and blade con- collar setae limbate, few in number. Side setae in
struction; side setae present in posterior thoracic third fascicle. Thoracic uncini with blunt, gouge-shaped
setigers from the third. pegs. Margin of collar fused dorsally. Incubation in
Paradexiospira Caullery and Mesnil 1897, Serpula faecal groove, attachment stalk present.
vitrea Fabricius 1780; 3 species. Simplicaria Knight-Jones, 1973, Spirorbis (Laeospira)
Dextrally coiled; three complete and one fourth pseudomilitaris Thiriot-Quievreux, 1965; 3 species.
incomplete thoracic setiger. Collar setae of fin and Sinistrally coiled; two pairs of thoracic tori present;
blade construction; side setae present in the third collar setae of fin and blade construction; blades usually
fascicle. Incubation in the tube. coarsely dentate, cross-striated. Side setae absent. In-
cubation in operculum.
Paralaeospira Caullery and Mesnil 1897, Spirorbis
(Paralaeospira) aggregata Caullery and Mesnil Spirorbella Chamberlin 1919, Spirorbis (Dexiospira)
1897; 8 species. marioni Caullery and Mesnil 1897; 3 species.
Sinistrally coiled; three thoracic tori on the concave Dextrally coiled; two pairs of thoracic tori; collar
side; collar setae of fin and blade construction without setae of fin and blade construction; side setae in third
cross-striations; side setae present in the third fascicle. fascicle. Thoracic uncini with broad anterior pegs. In-
Thoracic and abdominal uncini more numerous on the cubation in tube, attachment stalk absent.
concave side; thoracic uncini with blunt anterior pegs.
Spirorbis Daudin 1800, Spirorbis borealis Daudin
Incubation in faecal groove; attachment stalk absent.
1800; 48 species.
Pileolaria Claparede 1870a, P. militaris Claparede Sinistrally coiled; thorax with two pairs of tori; collar
1 870a; 20 species. setae of fin and blade construction; side setae present
Sinistrally coiled; two pairs of thoracic tori present; in the third fascicle. Thoracic uncini with broad an-
collar setae of fin and blade construction; blades usually terior pegs. Incubation in the tube, attachment stalk
coarsely dentate, cross-striated. Side setae present. absent.
Incubation in operculum.
Pillaiospira Knight-Jones 1973, Janua (Pillaospira) Taxonomic Notes

trifurcata Knight-Jones 1973; only species. Vine (1972) pointed out that the coiling of the spiror-
Dextrally coiled; two pairs of thoracic tori; collar bids may not be a character of particularly high value
setae limbate, abdominal setae with blades as long as and that even within one species, the coiling may vary
those in the collar setae; side setae absent. Incubation from sinistral to dextral. It is thus of the utmost im-
in the operculum. portance that great care be taken in using the key and
Prodexiospira Pillai 1970, Spirorbis violaceus Levinsen definitions given above and that all the important
1884; only species. features be checked carefully. It is quite possible that
Dextrally coiled; three complete and a fourth incom- the coiling will turn out to be of great value in most
plete setiger present. Collar setae limbate; side setae of the genera, even if the character should turn out
present in the third fascicle. Incubation in the tube. to be useless in certain instances.
Protolaeospira Pixell 1912, P. ambilateralis Pixell Invalid Genera

1912; 7 species.
Sinistrally coiled; three pairs of tort and in addition Charybs Montfort in Morch 1863, questionably Spi-
a fourth incomplete present. Collar setae of fin and rorbis
blade construction; side setae in third and fourth fas- Coretus Baster 1762, see Circeis
cicle. Thoracic uncini with bilobed anterior pegs. Incu- Dexiorbis Chamberlin 1919c, see Dexiospira
bation in faecal groove; attachment stalk present. Heterodisca Fleming 1825, indeterminable
Laeospira Caullery and Mesnil 1897, see Spirorbis
Protoleodora Pillai 1970, Spirorbis asperatus Bush Marsipospira Bailey 1969, see Protolaeospira
1904b; only species. Mera Saint-Joseph 1894, see Janua
Sinistrally coiled; three (two?) setigers in the thorax; Neodexiospira Pillai 1970, see Dexiospira

15 2 NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY Science Series 28
Pixellia Pillai 1970, see Protolaeospira consider them free-standing families of the Polychaeta.
Sinistrella Chamberlin 1919c, see Leodora Like Jouin and Westheide, I find it difficult to associate
Spirorbides Chamberlin 1919c, see Paradexiospira the five families, separately or as a unit, to any known
Stoa Serres 1855, indeterminable polychaete family, and have for that reason listed them
below, in alphabetical order. I do not think the five
FIVE "ARCHIANNELIDAN" FAMILIES families are particularly closely related to each other.
The following five families usually are considered

as belonging to a separate order, Archiannelida. They FAMILY DINOPHILIDAE REMANE 1932
are characterized, as pointed out by Hermans (1969),
by features adaptive to life in the mesopsammon, and Very small, usually distinctly segmented polychaetes,
the features in which they resemble each other are without appendages, setae and parapodia; some forms
exactly those that have been considered to be of selec- with adhesive glands posteriorly. Ciliary bands on
tive advantage for the mesopsammon (Swedmark 1964; ventral side present, and, usually segmentally arranged,
Hermans 1969). In agreement with Jouin (1971) and ciliary tracts present.
Westheide (1971), 1 find it difficult to retain the concept The family recently was reviewed by Jouin (1971)
of the Archiannelida (Fauchald 1974a) and presently and by Westheide (1971).
Key to Genera
Ia. Muscular pharynx absent Apharyngtus

l b. Muscular pharynx present 2
2a (1b). Body slender and elongated; prostomium trilobed Trilobodrilus
2b (Ib). Body short, more or less inflated; prostomium rounded 3
3a (2b). Pygidium with two adhesive glands; eyes absent Diurodrilus
3b (2b). Pygidium conical, without adhesive glands; eyes usually present Dinophilus
Generic Definitions on conical projections of various shapes. Pharyngeal

apparatus well developed.
Apharyngtus Westheide 1971, A. punicus Westheide
1971; only species. Trilobodrilus Remane 1925, T. heideri Remane 1925;
Slender, distinctly segmented bodies; pharyngeal 3 species.
apparatus absent; in addition to the regularly arranged Body long and slender; prostomium trilobed; eyes
ciliary bands, very fine cuticular hairs covering the absent; pygidium rounded or conical, without adhesive
body. glands. Pharyngeal apparatus well developed.
Dinophilus Schmidt 1848, D. vorticoides Schmidt FAMILY NERILLIDAE LEVINSEN 1883

1848; 8 species.
Interstitial small forms with few segments. Pros-
Short-bodied, indistinctly segment dinophilids; tomium with paired palps, nuchal organs and up to
pygidium rounded or conical. Pharyngeal apparatus
three antennae. Ventral eversible pharynx with four
present and well developed. Some forms distinctly jaws or unarmed. Uniramous parapodia with capillary
sexually dimorphic. setae or composite setae in a single fascicle. Para-
Diurodrilus Remane 1925, D. minimus Remane 1925; podial cirri usually present.
5 species. The key to genera and definitions given below have
Body short, inflated and indistinctly segmented. been modified after Jouin (1971). The main changes
Pygidium with paired adhesive glands ending distally involve terminology to fit the terms used in this paper.
Key to Genera
la. Body with nine segments including the post-peristomial pharyngeal segment 2
lb. Body with less than nine segments 4
2a (Ia). Composite setae present Mesonerilla
2b (Ia). Simple capillary setae present 3
3a (2b). Three antennae present Nerilla
3b (2b). Maximally one antenna present Meganerilla
f u
in
FIGURe 39. (A), Family DINOPHILIDAE, Diurodrilus ankeli, from Jouin, 1971, about 50x; (B), Family POLYGORDIIDAE,
Polygordius neapolitanus, after Jouin, 1971, about 15x; (C), posterior end of the above, about 15x; (D), Family SACCOCIRRIDAE,
Saccocirrus polycercus, after Jouin, 1971, about 25x; (E), posterior end of the above, about 25x; (F), Family PROTODRILIDAE,
Protodrilus hatscheki, after Jouin, 1971, about 15x; (G), posterior end of the above, about 15x; (H), Family NERILLIDAE,
Mesonerilla sp., after Jouin, 1971, about 25x.

4a (I b). Eight segments, including the post-peristomial pharyngeal segment 5

4b (1 b). Seven segments present 8
5a (4a). Both simple and composite setae present Nerillidopsis
5b (4a). All setae either composite or simple 6
6a (5a). All setae composite Thalassochaetus
6b (5a). All setae simple and capillary 7
7a (6b). Pharyngeal segment asetigerous, antennae absent; freshwater Troglochaetus
7b (6b). Pharyngeal segment with setae, two antennae present; marine Nerillidium
8a (4b). Composite setae; parapodial cirri absent, palps reduced Paranerilla
Sb (4b). Setae simple and capillary; parapodial cirri present except on last segment Psanunoriedlia
Generic Definitions Body with seven segments including the pharyngeal

segment. Antennae and parapodial cirri absent; palps
Meganerilla Boaden 1961, M. swedmarki Boaden
reduced to triangular lateral corners on the prostomium.
1961; 2 species.
Mud-dwelling forms. Setae composite.
Body with nine segments including the pharyngeal
segment. Two large palps and maximally one antenna Psammoriedlia Kirsteuer 1966, P. ruperti Kirsteuer
present. All setae simple capillaries. 1966; only species.
Mesonerilla Remane 1949, M. luederitzi Remane Body with seven segments including the pharyngeal
1949; 6 species. segment. Antennae absent; two palps present; para-
Body with nine segments including the pharyngeal podial cirri present, except on last segments. Setae
segment. Two palps and three antennae present. All simple capillaries.
setae composite. Thalassochaetus Ax 1954, T. palpifoliaceus Ax 1954;
Nerilla Schmidt 1848, N. antennata Schmidt 1848; only species.
7 species. Body with eight segments including the pharyngeal
Body with nine segments including the pharyngeal segment. Antennae and parapodial cirri absent; palps
segment. Three antennae and two palps present. All present. All setae composite spinigers.
setae simple capillaries.
Troglochaetus Delachaux 1921, T. beranecki Delachaux
Nerillidium Remane 1925, N. gracile Remane 1925; 1921; only species.
5 species. Body with eight segments, including the pharyngeal
Body with eight segments including the pharyngeal segment. Antennae absent; palps present; pharyngeal
segment. Pharyngeal segment with setae; maximally segment asetigerous. All setae simple capillaries.
two antennae present, may be absent. All setae simple Freshwater and caves.
capillaries.
Nerillidopsis Jouin 1967, N. hyalina Jouin 1967; only FAMILY POLYGORDIIDAE CZERNIAVSKY 1881a
species.
Body with eight segments including the pharyngeal
Slender, mainly interstitial forms. Prostomium with
segment. Anterior and posterior setae simple capillaries;
two solid antennae and nuchal slits. Eversible or non-
intermediate segments with composite setae. eversible muscular pharynx ventral. Eyes and parapodia
Paranerilla Jouin and Swedmark 1965, P. limicola absent; setae absent in most forms; segmentation and
Jouin and Swedmark 1965; only species. ciliation poorly developed.
Key to Genera
Ia. Setae present in posterior segments Chaetogordius

lb. Setae absent Polygordius
Generic Definitions Prostomium conical with paired, slightly articulated

antennae and large nuchal organs. Segmentation in-
Chaetogordius Moore 1904a, C. canaliculatus Moore distinct anteriorly, more distinct posteriorly. Single
1904a; only species. slender capillary setae in each of the last 10-12 seg-

ments; only one segment with paired setae on either FAMILY PROTODRILIDAE CZERNIAVSKY 1881a
side; all others with a single seta on either side.
Polygordius Schneider 1868, P. lacteus Schneider Interstitial slender forms. Prostomium with paired,
1868; 15 species. hollow or solid antennae; eyes present. Pharynx mus-
Prostomium rounded or conical with paired, usually cular, but not eversible. Segmentation and parapodia
smooth antennae and large nuchal organs. Segmenta- poorly developed, setae absent in most known species;
tion usually indistinct along the whole body. Posterior parapodia always absent.
end of body usually slightly wider than the rest of The family has been reviewed recently by Jouin
the body. Setae absent. (1966).
Key to Genera
Ia. Setae or segmentally arranged adhesive organs present Protodriloides
l b. Setae and segmentally arranged adhesive organs absent Protodrilus
Generic Definitions FAMILY SACCOCIRRIDAE CZERNIAVSKY 1881a

Protodriloides Jouin 1966, Protodrilus chaetifer Re-
mane 1926; 2 species. Interstitial slender forms. Prostomium with paired
Salivary glands and lateral organs absent; antennae hollow antennae; eyes present. Proboscis a ventral
solid. Setae or segmentally arranged adhesive glands muscular pad, present in most forms, but may be
present. absent. Parapodia uniramous, with simple, chisel-
Protodrilus Czerniavsky 1881a, P. mirabilis Czern- shaped setae.
iavsky 1881a; 20 species. The family is known for a single genus, Saccocirrus
Salivary glands and lateral organs present; antennae Bobretzky 1872, with genotype, S. papillocercus Bo-
hollow. Setae and segmentally arranged adhesive bretzky 1872 and about 12 species. Jouin (1971) gave
glands absent. a key to species.

GLOSSARY branchia (e)-any extension of the body wall with a

loop of the vascular system or which is well-
The glossary does not contain terms in general use equipped with capillary blood-vessels; a gill (27).
in zoology or in biology in general, such as distal and buccal-pertaining to the mouth.
proximal, anterior and posterior, etc. Generally, these
terms can be found in a good dictionary. The glossary
contains the terms characteristically used in systematic
polychaete literature and explanation of idiosyncratic capillary (ies)-in polychaete literature used either
usage where terms are used in a different sense from as an adjective in the combination capillary seta
the usual lexicographic one. Number in parenthesis or as a noun characterizing long, slender tapering
refer to the specific example of this feature illustrated setae.
in the appended illustrations. caruncle-posterior sensory organ projecting from the
prostomium sometimes over the first several
segments (28).
abdomen-posterior part of the body, behind the tho-
rax, sometimes followed by a tail (2). cephalon-head.
aciculum (a)-stout supportive setae found internally cephalic cage-long protective setae enclosing and
in each parapodial ramus where these project protecting the head.
from the body. Acicular setae are thick, project- cephalic rim-flange encircling the head (25).
ing setae (5).
cephalic veil-hoodlike membrane between the paleae
aileron-accessory jaw plate in the glycerids (9). and the buccal tentacles in pectinariids.
anal cirrus (i)-one or more elongated projections ceratophore-basal joint of an antenna (30).
from the pygidium (11).
ceratostyle-distal joint of an antenna (29).
antenna (e)-sensory projection arising from the
dorsal, lateral or frontal surface of the prostomium; chevron-V-shaped chitinized jaw piece at the base
innervated from the first part of the brain (12,61). of the eversible pharynx in some goniadids (31).
apodous-without parapodia (66). cirriform-slender and cylindrical (41).
aristate-about setae: simple setae with smooth shaft cirrophore-basal joint of a cirrus (35).
and a tuft of hairs or a single spine distally (17). cirrostyle-distal joint of a cirrus (34).
arborescent-branching (like a tree) (18). cirrus (i)-sensory projection, usually slender and
asetigerous-without setae (66). cylindrical, from the superior part of the noto-
podium (dorsal cirrus) or from the inferior part
auricular-ear-shaped (19). of the neuropodium (ventral cirrus) (36,38,78,80).
avicular-beaked (shaped like a bird's head) (20). clavate-club-shaped (32).
companion setae-small, simple setae in rows, ac-
biarticulate-with two joints; used about antennae, companying, or alternating with larger setae,
tentacles and palps (13). usually hooks of some kind.
bidentate-with two teeth (21). compound (or composite) setae-jointed setae (37).
bifid-split in two (22).

bifurcate-with two prongs (23). deciduous-liable to fall off (like leaves).
bilimbate-simple setae with two wings or flattened dentate-with teeth (39).

margins (24). denticulate-with small teeth (40).
bipinnate-formed like a feather with a main stem digitiform-finger-shaped (42).
and two rows of side branches (26).
biramous-with two branches; usually used about
parapodia having both noto- and neuropodia elytron (a)-dorsal scales found in the aphroditoids;
present (6). homologous with the dorsal cirrostyles (43).
elytrophore-aphroditoid cirrophore carrying an ely-

tron (44).
epitoke-modified reproductive stage or specimen,
often swarming.
facial tubercle-projecting ridge or lobe on the upper

lip of certain polychaetes (especially scale-
worms).
falcate-distally curved, usually blunt (used about
setae) (45).
falciger-distally blunt and curved setae (45).
filiform-slender and threadlike (46).
foliaceous-leaflike (47).
fusiform-cigar- or spindle-shaped (48).
geniculated-bent (like a knee) (49).
harpoon seta-stout pointed seta with recurved barbs

near the tip (50). A,
hemigomph articulation -with asymmetrical articu-

lation nearly at right angles to the long axis of
the shaft (52).
heterogomph articulation-with articulation clearly
oblique to the long axis of the shaft (53).
homogomph articulation-with articulation distinctly
and symmetrically at right angles to the long
axis of the shaft (54).
hooded setae-setae distally covered by a delicate
chitinous envelope or guard (51).
hook-general term used about stout-shafted, blunt,
often distally curved and dentate setae; smaller
hooks arranged in single or double rows are often
referred to as uncini.
imbricated-overlapping (like tiles).

inferior-the more ventral (of two or more structures).
ligule-finger-shaped major process on a parapodium

(55).
limbate-simple seta with a flattened margin or wing
(56).
lobe-major parapodial process, used mainly about
flattened kinds, but also more generally about all FIGURe 40. Diagram of selected terms defined in the Glossary
kinds of major parapodial processes (8). #1-18.
20
27
-50
-28
29 Pwl -52
mp
lo
11
Ficuan 41. Diagram of selected tens defined in the Glossary
#19-60.

1976 THE POLYCHAETE WORMs 159
long-handled-used about uncini, with a long basal

62 rod supporting the uncinus
63
I
metastomium-rarely used term covering the seg-
mented body between the prostomium and pygid-
ium, but including neither.
moniliform-beaded or beadlike (57).
mucro-with a sharp point or tip; abruptly tapered (58).
multiarticulated-with several joints (60).
natatory-swimming.
neuropodium (a)-ventral branch (ramus) of a para-
podium (7).
neurosetae-setae of the neuropodium.
notopodium (a)-dorsal branch (ramus) of a para-
podium (3).
notosetae-setae of a notopodium.
nuchal-pertaining to the neck; used about sensory
organs found on the posterodorsal side of the
head and variously developed as paired or single
processes, pits or grooves, sometimes as paired
epaulettes stretching posterolaterally from the
prostomium.
occipital-pertaining to the posterodorsal part of the

prostomium (64).
ocular-pertaining to the eyes.
ocular peduncle-projecting structure supporting the
eyes, especially in the polyodontids.
ommatophore-see ocular peduncle.
operculum (a)-literally a lid; used about a structure
some tubicolous worms use as stoppers for their
tubes when the occupant is retracted; see Sabell-
ariidae and Serpulidae.
palea (e)-strong or broad, usually flattened setae.

palmate-resembling the fronds of a palm (67).
I
palp (s)-sensory or feeding structures innervated
from the posterior part of the brain or from the
circumesophagal nerve ring; anteroventral and
sensory in the Phyllodocida; posterodorsal and
used in feeding in the spioniform worms (13).
FIGURE 42. Diagram of selected terms defined in the Glossary paragnath-chitinous denticle in the pharyngeal
#61-83. cavity of nereids (63).

parapodium (a)-segmentally arranged projections seta-secretion from the parapodia forming the arma-
carrying setae; foot (16). ture of these structures.
pectinate-comblike; with series of projections ar- setiger-segment carrying setae.
ranged like the teeth of a comb (68).
short-handled-used about uncini, without a long
penicillate-brushlike (70). rod-shaped support, fine threads may be present.
peristomium (a)-first distinct post-prostomial region; simple seta-unjointed seta.
strictly including only the region around the mouth,
in practice including also segments fused to this spatulate-blade-shaped, usually blunt-tipped, some-
structure, forming the posterior part of the recog- times with a mucro (59).
nizable head (16,66). spiniger-seta that tapers to a fine point; most fre-
pharynx (ges)-anterior part of the digestive tract; quently used about composite setae (33).
often eversible, always modified for feeding pur- spinous pocket-enlarged, pocketlike serration along
poses, sometimes also for burrowing (62). the margin of the setae of some scale-worms (74).
pinnate-featherlike, with a main stem and lateral stylode-small, fingerlike projection associated with
branches, see also bipinnate (71). a parapodium, usually small and distinctly longer
than wide (73).
plumose-resembling down; hairy (69).
postsetal-posterior to the setae; used about parapodial sub-biramous-pertaining to parapodia in which the
neuropodia are well developed and the notopodia
lobes or ligules (8).
reduced (77).
presetal-anterior to the setae; used about parapodial
l obes or ligules (4). subulate-awl-shaped; tapering to a fine point (75).
prostomium (a)-anteriormost, a pre-segmental part superior-the more dorsal (of two or more structures).
of the body anterior to the mouth, enclosing at
least the anterior part of the brain, often with
tentacular cirrus-sensory projections arising either
antennae and eyes (14,64).
from the peristomium or from cephalized seg-
proventricle-muscularized region of the anterior ments, in the latter case considered homologous
digestive tract in syllids, found posterior to the with the dorsal and ventral cirri of normal. prst-
pharynx. cephalic parapodia (15,65).
pygidium (a)-post-segmental terminal part of the thorax-anterior region of the body, Posterior to the
body carrying the anus (10). head (1).
trepan-chitinized, anteriorly toothed part of the
eversible pharynx of some worms, especially
radiole-one of the main tentacles in the tentacular
syllids (81).
crown of sabellids and serpulids.
ramose-branched. tridentate-with three teeth (82).
reniform-kidney-shaped (76).
rugose-roughened, lumpy (72). nations (i)-general term covering sharply dentate,
deeply imbedded setae, often with a platelike
base; or small, S-shaped setae with a distinct
beak. Uncini usually are arranged in rows trans-
scaphe-flattened caudal appendage of pectinariids.
verse to the long axis of the animal (83).
secondary tooth-the second of two teeth, the first
unciniger-segment carrying uncini.
being the apical, terminal or primary one.
unidentate-distally entire (45).
segment-any part of the body, apart from the pros-
tomium and pygidium set off internally or ex- uniramous-with one branch only; used about para-
ternally by septa or otherwise from the preceding podia in which one ramus, most frequently the
and following parts. notopodium, is absent (79).
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INDEX
Page numbers set in bold face type represent citations of illustrations.
A Amphitritides, 129, 130 Arenicolidae, 8, 13, 31, 37

Abarenicola, 37 Amphitritoides, 133 Arenicolides, 37
Aberranta, 25 Amphytrite, 133 Areniella, 35
Abyssoclymene, 39 Amplaria 149, 150 Arete, 91
Acanthicolepis, 58, 59 Amythas, 123, 125 Ariapithes, 120
Aceronereis, 91 Amythasides, 123, 125 Aricia, 17, 22
Acesta, 18 Amytis, 85 Aricidea, 17, 18
Acholoe, 59 Anaitides, 48 Aricideopsis, 25
Acicularia, 100 Anamobaea, 140 Arippasa, 140
Acoetes, 66 Anaspio, 23 Aristenia, 117
Acrocirridae, 8, 13, 14, 29, 30-31 Ancistargis, 78 Armandia, 42
Acrocirrus, 30 Ancistria, 35 Artacama, 128, 130
Admetella, 59 Ancistrosyllis, 78 Artacamella, 128, 130
Adyte, 57, 60 Andresia, 56, 60 Arwidssonia, 41
Aedicira, 17, 18 Andromache, Ill Aryandes, 128
Aenone, Ill Anguillosyllis, 80, 81 Asabellides, 124, 125
Aglaophamus, 97 Anisoceras, 113 Asclerocheilus, 44
Aglaopheme, 97 Anisocinus, 133 Asloegia, 103
Aglaura, 111 Anobothrella, 123, 125 Aspeira, 140
Aglaurides, I11 Anobothrus, 124, 125 Asterope, 52
Agnodice, 65 Anomalorbis, 148, 149, 150 Asychis, 39
Alcandra, 17 Anophthalmus, 77 Atelesyllis, 80, 81
Alciopa, 51, 52 Anoplonereis, 77 Athelepus, 133
Alciopidae, 8, 12, 46, 51-52 Anoplopisione, 73 Auchenoplax, 121, 125
Alciopina, 51, 52 Anoplosyllis, 85 Audouinia, 30
Alentia, 59, 60 Anotomastus, 33, 36 Augeneria, 109
Alentiana, 59, 60 Anthostoma, 17 Augeneriella, 137, 138
Alikunhia, 73, 75 Antinoana, 56, 60 Australaugeneria, 56, 60
Alitta, 91 Antince, 57, 60 Australonereis, 87, 88
Alkmaria, 123, 124 Antinoella, 57, 60 Austrolaenilla, 57, 60
Allia, 18 Antiobactrum, 42 Austrophyllum, 47, 48
Allmaniella, 58, 60 Antonbruunia, 91 Autolytides, 85
Alluaudella, 79, 81 Antonbmunidae, 8, 12, 91 Autolytus, 79, 81
Aloysina, 43 Aonides, 22, 23 Autosyllis, 79, 81
Amaea, 133 Aonis, 25, 97 Axionice, 130
Amaeana, 129, 130 Aonopsis, 25 Axiothea, 41
Amage, 124 Aotearia, 109 Axiothella, 39
Amblyosyllis, 81 Aparaonis, 17, 18
Ambo, 30 Apharyngtus, 152 B
Amelinna, 121, 124 Aphelothrix, 107
Americonuphis, 104, 105 Aphlebina, 133 Baffinia, 129, 131
Ammochares, 115 Aphrodita, 53, 54 Balanochaeta, 117
Ammotrypane, 43 Aphroditidae, 8, 10, 53-55, 54 Barantolla, 32, 33, 36
Ammotrypanella, 42 Aphrogenia, 54 Barbularia, 55
Ampharana, 123, 124 Apistobranchidae, 8, 14, 21, 22 Barrukia, 57, 60
Ampharete, 123, 124 Apistobranchus, 22 Bathya, 131, 133
Ampharetidae, 8, 10, 11, 121- Apneuma, 133 Bathyadmetella, 59, 60
128,122 Apomatopsis, 147 Bathychloeia, 100, 102
Ampharetidas, 134 Apomatus, 141, 143 Bathyglycinde, 92, 93, 94
Amphelothrix, 107 Aponobranchus, 135 Bathymoorea, 58, 60
Amphibranchus, 103 Apophryotrocha, 112 Bathynce, 65
Amphicora, 140 Apoprionospio, 23 Bebryce, 98
Amphicorina, 140 Aporosyllis, 85 Benhamipolynoe, 59, 61
Amphicteis, 124, 125 Aquilaspio, 23 Benthoscolex, 100, 102
Amphictene, 120 Arabella, 110, Ill Bergstroemia, 48, 49
Amphiduros, 75 Arabellidae, 8, 13, 108, 110-111 Berkeleya, 15, 16
Amphiglena, 137, 138 Arabes, Ill Bhawania, 72
Amphinome, 100, 102 Amcoda, 111 Biborin, 110
Amphinomidae, 8, 11, 100-103, Archidice, 30 Bilobaria, 25
101 Arcteobia, 57, 60 Bispira, 137, 138
Amphiro, 107, 133 Arcmnoe, 59, 60 Blainvillea, 107
Amphiron, 107 Arctonoella, 59, 60 Blends, 103
Amphisamytha, 124, 125 Arenia, 35 Boccardia, 22, 24, 25
Amphitrite, 129, 130 Arenicola, 37 Boguea, 135
Bogueidae, 8, 14, 122, 135 Chaetosphaera, 25, 65 Cryptonota, 104

Boguella, 135 Chaetosyllis, 85 Cryptopomatus, 120
Bonhourella, 142, 143 Chaetozone, 29 Ctenodrilidae, 8, 13, 19, 19-20
Bonuania, 74, 75 Chaponella, 41, 140 Ctenodrilus, 19, 20
Bookhoutia, 93 Charybs, 151 Ctenospio, 26
Bouchiria, 58, 61 Chaunorhynchus, 91 Cyanippa, 55
Brachysyllis, 85 Cheilonereis, 86, 88 Cyaxares, 133
Brada, 116 Cheironotus, 26 Cystonereis, 85
Bradabyssa, 116 Cherusca, 27 Cystopomatus, 147
Branchamphinome, 100, 102 Chinonereis, 91
Branchetus, 17 Chitinopoma, 143
D
Branchioasychis, 39 Chitinopomoides, 143, 144
Branchiocapitella, 32, 33, 36 Chloeia, 100, 102 Dalhousia, 77
Branchiomaldane, 37 Chloenea, 102, 103 Dalhousiella, 75, 76
Branchiomma, 137, 138, 140 Chloenopsis, 100, 102 Danymene, Ill
Branchionemis, 91 Chloochaeta, 103 Dasybranchetus, 33, 33, 36
Branchiosabella, 128 Chloraema, 117 Dasybranchus, 33, 33, 36
Branchiosyllis, 81 Chone, 137, 138 Dasychone, 140
Branchoscolex, 35 Chorizobranchus, 37 Dasychonopsis, 140
Brandtika, 140 Chrysodon, 120 Dasylepis, 65
Brania, 80, 82 Chrysopetalidae, 8, 10, 71, 71-72 Dasymallus, 36
Braniella, 80, 82 Chrysopetalum, 72 Dasynema, 142, 144
Briaraea, 99 Chrysothemis, 41 Dawbinia, 87, 88
Bruunilla, 118 Circeis, 149, 150 Decamastus, 32, 34, 36
Bucherta, 35 Cirraria, 51 Dejoces, 133
Bushiella, 149, 150 Cirratulidae, 8, 13, 28, 29-30 Demonax, 137, 138
Buskiella, 116 Cirratulispio, 30 Dendrobranchus, 133
Bylgia, 65 Cimatulus, 29 Dendronereides, 87, 88
Bylgides, 56, 61 Cirrhineris, 30 Dendronereis, 86, 88
Cirrifonnia, 29, 30 Dendrophora, 133
C Cirrobranchia, Ill Desdemona, 137, 138
Cirroceros, 91 Desmosyllis, 85
Cabira, 78 Cirrodoce, 47, 49 Dexiorbis, 151
Caesicirrus, 41 Cirronereis, 88, 91 Dexiospim, 148, 149, 150
Calamyzas, 85 Cirrophorus, 17, 18 Dextralia, 140, 150
Calamyzidae, 8, 14, 74, 85 Cirrosyllis, 77, 85 Dialychone, 137, 138
Calcareopomatus, 143 Cistena, 120 Didymobranchus, 103
Califia, 16 Cistenides, 120 Dilepidonotus, 58, 61
Callizona, 52 Claparedia, 85 Dindymene, 43
Callizonella, 52 Clavadoce, 48, 49 Dindymenides, 42
Campesyllis, 80, 82 Clavisyllis, 80, 82 Dinophilidae, 8, 9, 152, 153
Caobangia, 141 Clavodomm, 98 Dinophilus, 152
Caobangiidae, 8, 11, 136, 141 Cleta, 52 Diopatra, 104, 105
Capeospira, 149, 150 Clymaldane, 39, 40 Dioplosyllis, 80, 82
Capitella, 31, 32, 33, 36 Clymene, 41 Diplobranchus, 97
Capitellethus, 32, 33, 36 Clymenella, 39, 40 Diploceraea, 85
Capitellidae, 8, 13, 31, 31-36 Clymenides, 37 Diplocirrus, 116, 117
Capitellides, 32, 33, 36 Clymenopsis, 38, 40 Diplotis, 26
Capitita, 35 Clymenura, 39, 40 Dipolydora, 26
Capitobranchus, 33, 36 Clytie, 17 Dipomatus, 141, 144
Capitomastus, 31, 32, 33, 36 Coelobranchus, 110 Disoma, 26, 27
Carazzia, 25 Colobranchus, 26 Disomides, 27
Cambia, SI Colonianella, 103 Dispio, 23, 24
Cassandane, 43 Colymmatops, 129, 131 Distylia, 140
Castalia, 77 Commensodomm, 98 Distylidia, 137, 138
Caulleriella, 29 Conconia, 71 Ditrocha, 36
Cenogenus, 109 Conopomatus, 142, 144 Ditrupa, 142, 144
Cenothrix, III Coppingeria, 116, 117 Diurodrilus, 152
Centrocorone, 120 Corephoms, 135 Dodecaceria, 29, 30
Ceratocephale, 87, 88 Coretus, 151 Dorvillea, 112
Ceratonereis, 87, 88 Corynocephalus, 52 Dorvilleidae, 8, 11, 12, 13, 111-
Cervilia, 56, 61 Cossura, 21 113, 112
Chaetacanthus, 58, 61 Cossuridae, 8, 13, 20, 21 Doyeria, 85
Chaetobranchus, 133 Crithida, 85 Drieschella, 59, 61
Chaetogordius, 154 Crosslandiella, 142, 144 Drieschia, 58, 61
Chaetoparia, 48, 49 Crossostoma, 128 Drilognathus, 110
Chaetopteridae, 8, 11, 28, 28-29 Cmcigera, 142, 144 Drilonereis, 110, 111
Chaetopterus, 28 Cryptonereis, 87, 88 Duplicaria, 149, 150
Dybowscella, 140 Eunotomastus, 33, 34, 36 Frennia, 58, 61

Dysponetus, 72 Eunotophyllum, 51 Friedericiella, 73, 75
Eupallasia, 120
E Eupanthalis, 66 G
Euphione, 58, 61
Galathowenia, 114, 115
Ecamphicteis, 121, 125 Euphionella, 58, 61
Galeolaria, 142, 144
Echinorhynchus, 114 Eupholoe, 69, 70 Garjaiowella, 140
Eclysippe, 123, 125 Euphrosine, 103 Gastroceratella, 65
Egamella, 121, 125 Euphrosinidae, 8, 11, 101, 103 Gastrolepidia, 59, 61
Ehlersia, 81, 82 Eupista, 133 Gattiola, 85
Ehlersiella, 133 Eupistella, 129, 131
Gattyana, 57, 62
Ehlersileanira, 70 Eupolymnia, 130, 131
Geminosyllis, 81, 82
Eisigella, 36 Eupolynoe, 65
Genetyllis, 48, 49
Elicodasia, 27 Eupolyodontes, 66
Gisela, 17
Enapteris, 99 Eupomatus, 147
Globiodoce, 51
Enigma, 117 Eupompe, 66 Glossopsis, 147
Enipo, 57, 61 Euratella, 137, 138 Glycera, 91, 91
Enonella, 110 Eurato, 140
Glycerella, 91, 92
Enoplobranchus, 129, 131 Eurymedusa, 85
Glyceridae, 8, 11, 14, 91-92, 92
Eone, 94 Eurysyllis, 80, 82
(113)
Ephesia, 98 Eurythoe, 102
Glycinde, 92, 93, 94
Ephesiella, 98 Eusarnytha, 123, 125, 128 Glyphanostomum, 123, 125
Ephesiopsis, 98 Eusamythella, 123, 125 Glyphohesione, 79
Epicaste, 94 Euscione, 133 Gnatholycastis, 91
Epidiopatra, 104, 105 Eusclerocheilus, 45
Gnathosyllis, 85
Epimka, 53 Eusigalion, 71
Goniada, 93
Eracia, 51 Euspio, 26
Goniadella, 93
Eranno, 109 Eusthenelais, 71
Goniadidae, 8, 11, 14, 92-94, 93
Ereutho, 133 Eusyllis, 80, 82 (T13)
Eriographis, 140 Euthalenessa, 69, 70 Goniadides, 93
Eriphyle, 107 Euthelepus, 129, 131 Goniadopsis, 93
Escholtzia, 99 Euzonus, 42, 43
Gorbunovia, 140
Eteone, 47, 49 Evame, 65
Gorekia, 57, 62
Eteonella, 51 Evarnella, 65
Gossia, 85
Eteonides, 51 Exallopus, 112
Gravierella, 39, 40
Eteonopsis, 113 Exogone, 80, 82
Greeffia, 52
Ethocles, 22 Exogonella, 79, 82
Grubea, 85
Euarche, 66 Exogonita, 80, 82 Gmbeopolynoe, 58, 62
Eucarphus, 147 Exogonoides, 80, 82 Gmbeosyllis, 85
Eucarunculatus, 103 Exotokas, 85 Grubeulepis, 68
Eucerastes, 85 Gmbianella, 124, 125
Euchone, 41, 137, 138 F Grymaea, 133
Euchonella, 137, 138 Gunnarea, 118
Euclymene, 39, 40 Fabricia, 137, 139
Gwasitoa, 45
Eucranta, 57, 61 Fabriciella, 140
Gymnonereis, 87, 89
Eudistylia, 137, 138 Fabriciola, 137, 139
Gymnorhynchus, 91
Eudontosyllis, 80, 82 Fabrisabella, 137, 139 Gymnosoma, 140
Euglycera, 92 Falklandiella, 17 Gyptis, 75, 76
Eugrymaea, 133 Fauveldora, 149, 150
Eulaeospira, 148, 149, 150 Fauvelia, 80, 82 H
Eulagisca, 58, 61 Fauveliella, 73
Eulalia, 48, 49 Fauveliopsidae, 8, 13, 115, 117- Haematocleptes, 110
Eulalides, 51 118 Halimede, 77
Euleanhra, 71 Fauveliopsis, 118 Haliplanella, 53
Eulepethidae, 8, 10, 67, 68 Ficopomatus, 143, 144 Haliplanes, 53
Eulepethus, 68 Filibranchus, 134 Halithea, 55
Eulepis, 68 Filigrana, 147 Halla, l l l
Eumenia, 45, 51 Filograna, 142, 144 Halodora, 52
Eumeniopsis, 45 Filogranula, 142, 143, 144 Halogenia, 55
Eumida, 48, 49 Filopora, 147 Halosydna, 56, 59, 62
Eumidia, 51 Fimbriosthenelais, 69, 70 Halosydnella, 59, 62
Eumolpe, 65 Flabelliderma, 19, 116, 117 Halosydnoides, 65
Eunereis, 87, 88 Flabelligella, 30, 118 Halosydnopsis, 59, 62
Eunice, 106, 107 Flabelligera, 116, 117 Halyplanes, 53
Eunicidae, 8, 12, 105-107, 106 Flabelligeridae, 8, 10, 13, 115, Hamiglycem, 92
Euniphysa, 106 115-117 Haplobranchus, 140
Eunoe, 57, 61 Felmingia, 117 Haploscoloplos, 16
Eunomia, 51 Flora, 118 Haplosyllides, 81, 82
Haplosyllis, 81, 82 Hyboscolex, 44 Labotas, 17

Harmopsides, 65 Hydroides, 143, 144, 147 Labranda, 30
Hannothoe, 56, 57, 62 Hydrophanes, 53 Labrorostratus, 110
Harpochaeta, 79 Hypania, 124, 125 Lacharis, 94
Hartmania, 57, 62 Hypaniola, 124, 125 Lacydes, 17
Haswellia, 71 Hypephesia, 98 Lacydonia, 94
Hauchiella, 129, 131 Hypereteone, 51 Lacydoniidae, 8, 12, 94, 94
Hediste, 85, 88, 89 Hyperhalosydna, 59, 62 Ladice, 43
Hedyle, 91 Hypocirrus, 51 Laenilla, 65
Hekaterobranchus, 26 Hypoeulalia, 51 Laeonereis, 87, 89
Helena, 147 Hypsicomatides, 140 Laeospira, 151
Helicosiphon, 149, 150 Hypsicomatopsis, 140 Laetmatonice, 55
Hemilepidia, 57, 62 Hypsicomus, 137, 139 Laetmonice, 54
Heminereis, 91 Lagis, 120
Hemipodia, 92 I Lagisca, 57, 63
Hemipodus, 91, 92 Lais, 111
Ichthyotomidae, 8, 11, 113, 114
Hemisyllis, 85 Lalage, 85
[chthyotomus, 114
Heptaceras, 105 Lamellisyllis, 80, 83
Idalia, 134
Herdmanella, 56, 62 Lanassa, 129, 131
Idanthyrsus, 118
Hermadion, 57, 62 Langerhansia, 65, 85
Hermella, 120 Inermonephtys, 97 Lanice, 130, 131
Hermenia, 58, 62 llyphagus, 116, 117 Lanicides, 130, 131
Hertnione, 55 Intoshella, 57, 63 Laomedom, 85
Ioda, 85 Laonice, 23, 24
Hennionopsis, 54, 55
Hermodice, 102 Iospilidae, 8, 14, 95, 95-96 Laonome, 137, 139
lospilopsis, 95, 96
Hennonia, 55 Laonomedes, 140
lospilus, 95, 96
Hermundura, 79 Laphania, 129, 131
Iphianissa, 41
Hesiocaeca, 74, 76 Laphaniella, 134
Iphinereis, 91
Hesione, 75, 76 Lapithas, 85
Iphione, 56, 63
Hesionella, 74, 76 Laranda, Ill
Iphionella, 65
Hesionidae, 8, 12, 73-77, 74 Larymna, 111
Hesionides, 75, 76 Iphitime, 109 Leaena, 129, 131
Hesionura, 47, 49 Iphitimidae, 8, 11, 108, 109-110 Leanim, 69, 70
Hesiospina, 75, 76 Iran, 121, 126 Ledon, 31
Hesiosyllis, 73, 76, 84 Irmula, 80, 83, 84 Leiobranchus, 140
Hesperalia, 85 Isocirrus, 39, 40 Leiocapitella, 33, 34, 36
Hesperonoe, 57, 62 Isolda, 121, 126, 127 Leiocapitellides, 32, 34, 36
Hesperophyllum, 47, 49 Isomastus, 36 Leiocephalus, 41
Heteraphrodita, 53, 54 Isosyllis, 85 Leiochone, 41
Heterobranchus, 128 Leiochrides, 32, 34, 36
J
Heterocirrus, 30 Leiochms, 32, 34, 36
Heteroclymene, 39, 40 Janita, 143, 144 Leipoceras, 26
Heterodisca, 151 Janua, 149, 150 Lenora, 103
Heteromaldane, 41 Jasmineva, 137, 139 Leocrates, 75, 76
Heteromarphysa, 107 Johnstonella, 99 Leocratides, 75, 76
Heteromastides, 31, 34, 36 Johnstonia, 39, 40 Leodice, 107
Heteromastus, 32, 34, 36 Josephella, 141, 144 Leodora, 148, 149, 150
Hetemnereis, 91 Leonnates, 87, 89
Heteropale, 72 K Leonnatus, 94
Heterophenacia, 133 Leontis, 91
Hererophyselia, 134 Kainonereis, 87, 89 Lepadorhynchus, 51
Heteropodarke, 75, 76 Kalaminochaeta, 27 Lepidametria, 65
Heteropolynoe, 57, 62 Kalaminochaetidae, 27 Lepidasthenia, 59, 63
Kalummaria, 27
Heterospio, 26, 27 Lepidastheniella, 59, 63
Kebuita, 44
Heterospionidae, 8, 14, 27-28, 29 Lepidia, 66
Kefersteinia, 75, 76
Heterosyllis, 85 Lepidofimbria, 58, 63
Kermadecella, 57, 63
Heteroterebella, 134 Lepidogyra, 58, 63
Kesun, 42, 43
Hipponoa, 100, 102 Lepidonereis, 91
Kinbergella, 26
Histriobdella, 114 Lepidonotus, 56, 58, 63
Kinbergja, 51
Histriobdellidae, 8, 12, 113, 113- Lepidopleurus, 71
Kinberginereis, 87, 89
114 Lepraea, 134
Histriodrilus, 114 Krohnia, 51, 52 Leptochaetoptems, 29
Kuwaita, 107, 109
Hololepida, 59, 62 Leptochone, 140
Hololepidella, 58, 62 Kynephorus, 79 Leptoecia, 105
Horstileanira, 69, 70 Leptonereis, 87, 89
L
Hyalinoecia, 104, 105 Letmonicella, 55
Hyalopomatopsis, 147 Labiaria, 121 Leucariste, 134
Hyalopomatus, 143, 144 Labidognathus, I11 Leucia, 57, 63

Leucodora, 26 Mandane, 26 Monticellina, 30

Levidorum, 97, 98 Mandrocles, 41 Morants, 22, 24
Levinsenia, 18 Mania, 77 Moyanus, 121, 126
Linophems, 100, 102 Marenzelleria, 23, 24 Mugga, 123, 126
Liocapa, 52 Marifugia, 142, 145 Muggoides, 123, 126
Lipephila, 91 Marphysa, 106, 107 Myriana, 51
Lipobranchius, 43, 44 Marsipospim, 151 Myrianida, 79, 83
Lirione, 103 Mastigethus, 53 Myriacyclum, 51
Lithognatha, 107 Mastigonereis, 91 Myriochele, 115
Litocorsa, 78 Mastobranchus, 32, 34, 36 Myrioglobula, 115
Loandalia, 78 Mauita, 52 Myriowenia, 114, 115
Loimia, 130,131 Maupasia, 53 Mysta, 47, 49
Longosoma, 28 Mayella, 69, 70 Mystides, 47, 49
Lopadorhynchidae, 8, 12, 46, 52- Mayeria, 107 Myxicola, 136, 139
53 Mediomastus, 32, 34, 36
Lopadorhynchus, 53 Megachone, 140
N
Lophocephalus, 117 Megalomma, 137, 139
Lophonota, 103 Meganereis, 91 Naiades, 51, 52
Lophosyllis, 85 Meganerilla, 152, 154 Naidonereis, 17
Lucopia, 58, 63 Meiodorvillea, 112 Nainereis, 15, 16
Lugia, 47, 49 Melaenis, 57, 63 Nais, 43
Lumara, 134 Melinantipoda, 121, 126 Namalycastis, 87, 89
Lumbriclymene, 38, 40 Melinella, 130, 131 Namanereis, 87, 89
Lumbriclymenella, 38, 40 Melinna, 121, 126 Naneva, 130, 131
Lumbricomastas, 33, 34, 36 Melinnampharete, 123, 126 Naraganseta, 30
Lumbriconais, 36 Melinnata, 123, 126 Naumachius, 91
Lumbriconereis, 109 Melinnexis, 121, 126 Nauphanta, 52, 107
Lumbrineridae, 8, 13, 107-109, 108 Melinnides, 128 Nausicaa, 107
Lumbrinerides, 109 Melinnoides, 123, 126 Neanthes, 86, 88, 89
Lumbrineriopsis, 109 Melinnopsides, 121, 126 Neco, 41
Lumbrineris, 107, 109 Melinnopsis, 121, 126, 128 Nectochaeta, 65
Lycaretus, 103 Membranopsis, 145, 147 Nectoneanthes, 88, 89
Lycastella, 91 Mera, 151 Nectonereis, 91
Lycastis, 85, 91 Mercierella, 143, 145 Nematonereis, 106, 107
Lycastoides, 87, 89, 91 Mercierellopsis, 143, 145 Nemidia, 57, 64
Lycastopsis, 87, 89 Mesochaetopterus, 28 Neoamphitrite, 129, 131
Lycidice, 107 Mesocirrineris, 30 Neodexiospira, 151
Lycoris, 91 Mesoeulalia, 51 Neoheteromastus, 32, 34, 36
Lygdamis, 118 Mesomystides, 51 Neohololepidella, 58, 64
Lysarete, Ill Mesonerilla, 152, 154 Neoleanira, 69, 70
Lysaretidae, 8, 11, 108, 111 Mesospio, 23, 24 Neoleprea, 129, 132
Lysidice, 106 Mesotrocha, 29 Neolipobranchius, 43, 44
Lysilla, 129, 131 Metachone, 140 Neomediomastus, 32, 34, 36
Lysippe, 124, 126 Metalaeospira, 149, 150 Neomeris, 43
Lysippides, 124, 126 Metalaonome, 140 Neomicrorbis, 148, 149, 151
Metamphinome, 103 Neonotomastus, 32, 34, 36
Metavermilia, 142, 145 Neopaiwa, 123, 126
M
Mexamage, 124, 126 Neopanthalis, 66
Macduffia, 107 Mexieulepis, 68 Neopodarke, 75, 76
Macellicephala, 56, 63 Microclymene, 39, 40 Neopomatus, 143, 145
Macellicephaloides, 56, 63 Micromaldane, 38, 40 Neopygospio, 26
Macelloides, 56, 63 Micronephthys, 97 Neosabellides, 123, 127
Maclovia, Ill Micronereides, 87, 89 Neosamytha, 123, 127
Macrochaeta, 30, 31 Micronereis, 87, 89 Neottis, 134
Macroclymene, 39, 40 Microphthalmus, 75, 76 Neovermilia, 143, 145
Macroclymenella, 39, 40 Micropodarke, 75, 76 Nephtyidae, 8, 12, 95, 96, 96-97
Macrophyllum, 51 Microrbinia, 15, 16 (T13)
Maea, 26 Microsamytha, 128 Nephtys, 97
Magelona, 26 (TIO) Microserpula, 147 Nereidae, 8, 12, 85-91, 86
Magelonidae, 8, 14, 26, 26 Microspio, 23, 24 Nereidice, 107
Malacoceros, 23, 24 Microsyllis, 85 Nereidonta, 107
Maldane, 39, 40 Milnesia, 55 Nereilepas, 91
Maldanella, 39, 40 Minuspio, 23, 24 Nereimyra, 75, 76
Maldanidae, 8, 13, 37-41, 38 Mitraria, 115 Nereiphylla, 48, 49
Maldanopsis, 41 Monocerina, 85 Nereis, 86, 88, 90
Malmgrenia, 59, 63 Monorchos, 118, 119, 120 Nereisyllis, 85
Malmgreniella, 59, 63 Monostylos, 20 Nereitube, 105
Manayunkia, 137, 139, 140 Monroika, 137, 139, 140 Nerilla, 152, 154
Nerillidae, 8, 12, 152-154, 153 Orbiniella, 15, 16 Parapolynoe, 65

Nerillidium, 154 Orbiniidae, 8, 13, 14-17, 15 Paraprionospio, 23, 24
Nerillidopsis, 154 Oria, 140 Parapterosyllis, 81, 83
Nerine, 26 Oriades, 140 Pamsabella, 140
Nerinides, 26 Oridia, 140 Parasclerocheilus, 44
Nerinopsis, 26 Oriopsis, 137, 139 Paraserpula, 143, 145
Nevaya, 27 Orseis, 73, 76 Parasphaerosyllis, 81, 83
Nicolea, 130, 132 Orthodromus, 77 Parasitosyllis, 85
Nicomache, 38, 40 Otanes, 134 Parasyllidea, 75, 77
Nicomachella, 41 Othonia, 140 Parathelepus, 129, 132
Nicomedes, 91 Otopsis, 78 Paratyposyllis, 81, 83
Nicon, 87, 90 Owenia, 114, 115 Parautolytus, 80, 83
Nicotia, 85 Oweniidae, 8, 11, 14, 114-115, 115 Paravermilia, 147
Ninoe, 107, 109 Oxydromus, 77 Paraxionice, 130, 132
Nipponophyllum, 47, 50 Paraxiothea, 41
Nitetis, 43 P Parergodrilidae, 8, 13, 19, 20
Norepa, 65 Parergodrilus, 20
Northia, 105 Pabits, 123, 127 Pareulepis, 68
Nossis, 91 Paedophylax, 85 Pareupholoe, 69, 70
Notalia, 48, 50 Pagenstecheria, 85 Pareurythoe, 100, 102
Notaulax, 140 Paiwa, 124, 127 Parexogone, 85
Nothis, 51 Paleanotus, 72 Parheteromastides, 32, 35, 36
Nothria, 105 Pallasia, 120 Parheteromastus, 32, 35, 36
Notocirrus, I10, 111 Pallasina, 120 Parhypania, 124, 127
Notodasus, 32, 34, 36 Pallonia, 134 Parmensis, 65
Notomastus, 32, 34, 36 Palmyra, 72 Paronuphis, 105
Notophyllum, 47, 50 Palmyreuphrosyne, 103 Parophryotrocha, 112
Notoproctus, 38, 41 Palmyridae, 8, 10, 71, 71, 72 Parthenope, 20
Notopsilus, 110, 111 Palola, 106, 107 Paumotella, 142, 145
Notopygos, 100, 102 Palpiglossus, 107 Pectinaria, 120
Novaquesta, 19 Panthalis, 66 Pectinariidae, 8, 10, 119, 120-121
Novobranchus, 134 Panousea, 138, 139 Pedinosoma, 53
Nuchubranchiata, 100 Pantoithrix, 116, 117 Peisidice, 68
Nudisyllis, 80, 83 Papillaria, 26 Peisidicidae, 66
Nychia, 65 Parachloeia, 100, 102 Pelagobia, 52, 53
Parachonia, 140 Pellucidaria, 97
Paradexiospira, 149, 151 Pelogenia, 71
0
Paradiopatra, 105 Pentacirms, 30
Octobranchus, 134 Paradoneis, 18 Peresiella, 31, 35, 36
Odontosyllis, 81, 83 Paradyte, 57, 64 Perialla, 26
Odysseus, 134 Paraeulalia, 51 Periboea, 73, 77, 85
Oenone, 111 Parahalosydna, 58, 64 Perinereis, 88, 90
Oeorpata, 127 Parahesione, 75, 77 Perouepis, 59, 64
Olga, 142, 145 Parahololepidella, 58, 64 Petaloclymene, 41
Olgaharmania, 143, 145 Paraiospilus, 95, 96 Petaloproctus, 39, 41
Oligobranchus, 45 Paralacydonia, 94 Petitia, 80, 83
Oligognathus, 110 Paralaeospira, 149, 151 Petta, 120, 121
Oligolepis, 65 Paralanice, 130, 132 Phaetusa, 91
Omaria, 43 Paraleiocapitella, 32, 35, 36 Phalacrophorus, 95, 96
Omphalopoma, 143, 145 Paralepidonotus, 59, 64 Phalacrostemma, 118, 119
Omphalopomopsis, 142, 145 Paralycastis, 91 Pharyngeovalvata, 80, 83
Oncoscolex, 45 Paramage, 124, 127 Phenacia, 134
Oniscosoma, 104 Paramarphysa, 106, 107 Pherea, 134
Onuphidae, 8, 10, 12, 104, 104-105 Paramphicteis, 124, 127 Pherecardia, 102, 103
Onuphis, 105 Paramphinome, 100, 102 Pherecardites, 102, 103
Oophylax, 85 Paranaitis, 48, 50 Phemsa, 116, 117
Ophelia, 42, 43 Parandalia, 78, 79 Phisidia, 129, 132
Opheliidae, 8, 14, 41-43, 42 Perinereis, 91 Pholoe, 69, 70
Ophelina, 42, 43 Paranerilla, 154 Pholoides, 66
Ophiodromus, 75, 76, 77 Paranerine, 26 Pholoididae, 8, 10, 66, 67
Ophioglycera, 93, 94 Paranorthia, 104, 105 Photocharis, 85
Ophiuricola, 107, 109 Paranychia, 65 Phragmatopoma, 118, 119
Ophryotrocha, 112 Paraonella, 18 Phronia, 79
Opisthodonta, 80, 83 Paraonidae, 8, 13, 14, 15, 17-18 Phyllampharete, 123, 127
Opisthopista, 129, 132 Paraonides, 18 Phyllamphicteis, 124, 127
Opisthosyllis, 81, 83 Paraonis, 18 Phyllochaetopterus, 28
Ops, 115 Parapholoe, 68 Phyllocomus, 124, 127
Orbinia, 16 Parapionosyllis, 80, 83 Phyllodoce, 48, 50, 51
Phyllodocidae, 8, 12, 45-51, 46 Pomatoceros, 142, 146 Pseudocapitella, 33, 35, 36

(T-13) Pomatoleios, 142,146 Pseudochitinopoma, 143, 146
Phyllohartmania, 57, 64 Pomatostegus, 142, 146 Pseudocirratulus, 29, 30
Phyllonereis, 91 Pontodora, 53 Pseudoclymene, 39, 41
Phyllosheila, 57, 64 Pontodoridae, 8, 12, 46, 53 Pseudofabricia, 137, 140
Phyllosyllis, 79, 83 Pontogenia, 54 Pseudohalosydna, 59, 64
Phylo, 16 Porroa, 51 Pseudolacydonia, 94
Physalidonotus, 65 Portelia, 97 Pseudoleiocapitella, 32, 35, 36
Phyzelia, 134 Potamethus, 138,139 Pseudoleucodore, 26
Pilargiidae, 8, 12, 74, 77-79 Potamilla, 138, 139, 140 Pseudomalacoceros, 23, 24
Pilargis, 78 Potamis, 140 Pseudonemis, 88, 90
Pilearia, 27 Potaspina, 138, 139 Pseudonerine, 26
Pileolaria, 149, 151 Pottsiscalisetosus, 58, 64 Pseudonotophyllum, 51
Pillaiospim, 148, 149, 151 Praegeria, 73 Pseudopalolo, 107
Pionosyllis, 80, 83 Praxilla, 41 Pseudopolydora, 22, 25
Pirakia, 48, 50 Praxillella, 39, 41 Pseudopolynoe, 58, 65
Piratesa, 147 Praxillura, 38, 41 Pseudopomatoceros, 142, 146
Piromis, 116, 117 Praxithea, 91 Pseudopotamilla, 140, 141
Pisenoe, 91 Prionognathus, 113 Pseudosabellides, 128
Pisione, 73 Prionospio, 22, 23, 24 Pseudoscalibregma, 44
Pisionella, 73 Proboscidea, 92 Pseudoserpula, 141, 146
Pisionidae, 8, 10, 12, 72, 72-73 Proceraea, 79, 83 Pseudosyllides, 81, 84, 85
Pisionidens, 73 Procerastea, 79, 84 Pseudosyllidia, 77
Pisionura, 72 Prochaetoparia, 48, 50 Pseudothelepus, 133
Pista, 130, 132, (T10) Proclea, 129, 132 Pseudovetmilia, 143, 146
Pixellia, 152 Proclymene, 39, 41 Psymobranchus, 147
Placostegus, 142, 145 Procome, 85 Pterampharete, 123, 127
Placostegopsis, 147 Prodexiospira, 149, 151 Ptemutolytus, 85
Plakosyllis, 80, 83 Progoniada, 92, 94 Pteriptyches, 26
Platynereis, 86, 88, 90 Promaldane, 41 Ptemcirrus, 48, 50
Platysyllis, 85 Promenia, 30 Pterolysippe, 124, 127
Pleione, 103 Prophyllodoce, 48, 50 Pteroscolex, 37
Plioceras, 111 Proplacostegus, 147 Pterosyllis, 85
Plotobia, 100 Proscalibregma, 43, 44 Pterothrix, 111
Plomhelmis, 51, 52 Proscoloplos, 15, 16 Pulliella, 32, 35, 36
Plotolepis, 65 Prospio, 26 Puparia, 37
Podarke, 75, 77 Protis, 141, 146 Pycnoderma, 117
Podarkeopsis, 75, 77 Protoaricia, 15, 16 Pygophelia, 43
Podarmus, 58, 64 Prdtoariciella, 15, 17 Pygophyllum, 26
Podioceros, 147 Protocapitella, 37 Pygospio, 23, 25
Podonereis, 85, 91 Protocarobia, 51
Poecilochaetidae, 8, 14, 27, 27 Protodorvillea, 112 Q
Poecilochaetus, 27 Protodrilidae, 8, 9, 12, 153, 155
Questa, 19
Poeobiidae, 8, 9, 115, 117 Protodriloides, 155
Questidae, 8, 13, 15, 18-19
Poeobius, 117 Protodrilus, 155
Quetieria, 65
Pollicita, 98 Protogrubea, 85
Polybostrichus, 85 Protolaeospim, 149, 151 R
Polybranchia, 22, 24 Protoleodora, 149, 151
Polycimrs, 129, 132 Protolycoris, 91 Ramex, 130, 132
Polydora, 22, 24, 25 Protomasmbranchus, 33, 35, 36 Ranzania, 29
Polydorella, 22, 24 Protomystides, 48, 50 Ranzanides, 29
Polyeunoa, 58, 64 Protopolydom, 26 Raphidrilus, 20
Polygordiidae, 8, 153, 154-155 Protothelepus, 134 Raricirrus, 29, 30
Polygordius, 154, 155 Protula, 146 Rashgua, 32, 35, 36
Polylepis, 71 Protulides, 140 Reibischia, 53
Polymastus, 85 Psammocollus, 115 Restio, 66
Polymnia, 134 Psammodrilidae, 8, 13, 20, 20-21 Reteterebella, 130, 132
Polymniella, 129, 132 Psammodriloides, 21 Rhamphobrachium, 104, 105
Polynice, 85 Psammodrilus, 21 Rhodine, 38, 41
Polynoe, 57, 64 Psammolyce, 69, 70 Rhodopsis, 142, 146
Polynoella, 56, 64 Psammoriedlia, 154 Rhopalosyllis, 80, 84
Polynoidae, 8, 10, 55, 55-65 Psectra, 72 Rhynchobolus, 92
Polyodonms, 66 Pseudampharete, 128, 132, 133 Rhynchonerella, 51, 52
Polyodontidae, 8, 10, 55, 65-66 Pseudeulalia, 48, 50 Rhynchospio, 23, 25
Polyophthalmus, 42, 43 Pseudeupanthalis, 66, 71 Rhynophylla, 26
Polyphragma, 147 Pseudeurythoe, 103 Robertianella, 56, 65
Polyphysia, 43, 44 Pseudexogone, 80, 84 Romanchella, 149, 151
Pomatoceropsis, 147 Pseudobranchiomma, 137, 140 Rostraria, 103
Rullierinereis, 87, 90 Sige, 48, 50 Subprotula, 141, 147

Rytocephalus, 128 Siliquaria, 147 Syllia, 85
Simplicaria, 148, 149, 151 Syllidae, 8, 12, 74, 79-85
S Sinistrella, 152 Syllides, 80, 84
Siphonostoma, 117 Syllidia, 75, 77
Sabaco, 41 Siphostoma, 117 Sylline, 85
Sabella, 137, 140 Sitophaga, 141 Syllis, 81, 84
Sabellaria, 118, 119, 120 Skardaria, 22 Synelmis, 78
Sabellariidae, 8, 10, 118-120, 119 Solomononereis, 87, 90 Synsyllis, 80, 84
Sabellastarte, 137, 140 Solowetia, 134
Sabellidae, 8, 11, 135-141, 136 Sonatsa, 39, 41 T
Sabellides, 123, 127 Some, 124, 128
Sabellina, 140 Sosanella, 123, 128 Tachytrypane, 42, 43
Sabellonga, 141 Sosanides, 124, 128 Tainokia, III
Sabellongidae, 8, 13, 136, 141 Sosanopsis, 124, 128 Talehsapia, 78
Sabidius, 18 Spermosyllis, 80, 84 Taphus, 72
Sabina, 140 Sphaerephesia, 97, 98 Tauberia, 18
Saccocirtidae, 8, 153, 155 Sphaerodoce, 48, 50 Tecturella, 117
Saccocirrus, 155 Sphaerodoridae, 8, 12, 95, 96, 97- Telake, 92
Sacconereis, 85 98 Telamone, 77
Sagitella, 99 Sphaerodoridium, 98 Telepsavus, 29
Salmacina, 141, 146 Sphaerodoropsis, 98 Telethusae, 37
Salmacinopsis, 141, 146 Sphaerodorum, 98 Telolepidasthenia, 59, 65
Salvatoria, 85 Sphaeropomatus, 143, 147 Telonereis, 113
Samytha, 123, 127 Sphaerosyllis, 80, 84, 97 Telothelepus, 129, 133
Samythella, 123, 127 Spinosphaera, 129, 132 Temporaria, 142, 147
Samythopsis, 123, 128 Spinther, 100, 104 Tenonia, 57, 65
Sandanis, 36 Spintheridae, 8, 9, 100, 101, 104 Terebella, 129, 133
Sangiria, 100, 103 Spin, 23, 25 Terebellanice, 130, 133
Schhobranchia, 117 Spiochaetopterus, 28 Terebellidae, 8, 11, 122, 128-134
Scalibregma, 44 Spione, 26 Terebellides, 134
Scalibregmella, 44, 45 Spionereis, 26 Terebellobranchia, 129, 133
Scalibregmidae, 8, 14, 42, 43-45 Spionidae, 8, 14, 21, 22-26 Terpsichore, 43
Scalibregmides, 44, 45 Spionides, 26 Tetraglene, 85
Scalis, 121 Spiophanes, 22, 25 Tetratrocha, 91
Scalisetosus, 57, 65 Spiramella, 147 Tetreres, 120
Scalispinigera, 43, 44, 45 Spirobranchus, 142, 147 Thalassema, 114
Schistocomus, 124, 128 Spirodiscus, 142, 147 Thalassochaetus, 154
Schistomeringos, 112, 113 Spirographis, 137, 140 Thalenessa, 69, 70
Schizobranchia, 137, 140 Spirorbella, 148, 149, 151 Tharyx, 29, 30
Schiaocraspedon, 142, 146 Spirorbidae, 8, 11, 147-152, 148 Thaumastoma, 27
Schmardanella, 134 Spirorbides, 152 Thelepella, 134
Schmardia, 85 Spirorbis, 149, 151 Thelephusa, 134
Schmardiella, 77 Spirovenna, 129, 133 Thelepides, 130, 133, 134
Schreiberius, 114 Staurocephalus, 113 Thelepodopsis, 134
Schroederella, 15, 17 Stauroceps, 113 Thelepus, 129, 133
Scione, 134 Stauronereis, 113 Theodisca, 17
Scionella, 130, 132 Steggoa, 48, 50 Theostoma, 17
Scionides, 130, 132 Steninonereis, 87, 90 Therochaeta, 116, 117
Scionopsis, 134 Stephania, 77 Therochaetella, 116, 117
Sclerobregma, 44, 45 Stephanosyllis, 85 Thesmia, 103
Sclerocheilus, 44, 45 Stemaspidae, 8, 10, 113, 114 Thetisella, 103
Sclerostyla, 142, 143, 146 Stemaspis, 114 Thoe, 85
Scolecolepides, 23, 25 Sthenelais, 70 Thoosa, 91
Scolecolepis, 26 Sthenelanella, 69, 70 Thormora, 58, 65
Scolelepis, 23, 25 Sthenolepis, 70 Thylaciphorus, 85
Scoletoma, 109 Stoa, 152 Thysanoplea, 98
Scoloplella, 15, 17 Strategis, 103 Tibiana, 107
Scoloplos (Leodamas) 16, 17 Stratiodrilus, 114 Timarete, 29, 30
Scoloplos (Scoloplos) 16, 17 Stratonice, 91 Tomopteridae, 8, 11, 95, 96, 98-99
Scyphoproctus, 32, 35, 36 Streblosoma, 129, 133 Tomopteris, 99
Semiodem, 117 Streblospio, 23, 25 Torquea, 134
Serpula, 143, 147 Streptosyllis, 80, 84 Torrea, 51, 52
Serpulidae, 8, 10, 11, 141-147, 148 Struwela, 74, 77 Trachelophyllum, 51
Sheila, 58, 65 Stschapovella, 129, 133 Trapodia, 105
Sigalion, 69, 70 Stygocapitella, 20 Travisia, 42, 43
Sigalionidae, 8, 10, 68-71, 69 Stylarioides, 117 Travisiopsis, 99
Sigambra, 78 Subadyte, 57, 65 Tricertia, 53, 54

Trichobmnchidae, 8, II, 122, 134- U Vitiaziphyllum, 48, 51

1 35
Trichobranchus, 134, 135 Umbellisyllis, 79, 84 w
Trichosobranchella, 140, 141 Unanereis, 87, 90
Trichosyllis, 85 Uncinereis, 91 Wartelia, 134
Tricoelia, 29 Uncinia, 26 Watelio, 51, 52
Tricosmochaeta, 65 Unciniseta, 109 Weberia, 59, 65
Trilobodrilus, 152 Uncinochaeta, 134 Websterinereis, 87, 90
Tripolydora, 22, 25 Uncopolynoe, 56, 65 Weddellia, 123, 128
Trochochaeta, 26 Unobranchus, 134, 135 Wesenbergia, 75, 77
Trochochaetidae, 8, 14, 21, 26-27 Urosiphon, 43 Willeysthenelais, 69, 70
Troglochaetus, 154
X
Trophonia, 117 V
Trophoniella, 116, 117 Xenosyllides, 85
Trypanosyllis, 79, 81, 84 Valla, 36 Xenosyllis, 81, 84
Tylonereis, 87, 90 Vanadis, 51, 52
Tylorrhynchus, 87, 90 Veleda, 103 z
Typhlonereis, 91 Venadis, 17 Zeppelinia, 20
Typhloscolecidae, 8, 14, 95, 99, Venusia, 134 Zopyrus, 147
99-100 Vennilia, 147 Zoms, 117
Typhloscolex, 99 Vetmiliopsis, 143, 147 Zothea, 103
Typosyllis, 81, 84 Virchowia, 85 Zygolobus, 109
Tyrrhena, 77 Vitiazia, 48, 50 Zygophyllus, 109

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THE POLYCHAETE WORMS

DEFINITIONS AND KEYS TO THE ORDERS, FAMILIES AND GENERA

NATURAL HISTORY MUSEUM

CHARACTERS USED TO DEFINE HIGHER TAXA 2

ORDER STERNASPIDA 114

ORDER OWENIIDA 114

ORDER FAUVELIOPSIDA 117

ORDER TEREBELLIDA 118

ORDER SABELLIDA 135

FIVE "ARCHIANNELIDAN" FAMILIES 152

T occupation, practised calmly and slowly, and

T gave during a class in polychaete biology at

INTRODUCTION and the setae, if present, only rarely occur in bundles.

2 NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY Science Series 28

1976 THE POLYCHAETE WORMS 3

4 NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY Science Series 28

1976 THE POLYCHAETE WORMS 5

6 NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY Science Series 28

1976 THE POLYCHAETE WORMS 7

8 NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY Science Series 28

Survey of Polychaete Orders, Suborders, Superfamilies and Families.

l. O. ORBINIIDA 7. O. OPHELIIDA SO. Glyceriformia 12. O. STERNASPIDA

1976 THE POLYCHAETE WORMS 9

Ia. External segmentation and setae absent 2

10 NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY Science Series 28

1976 THE POLYCHAETE WORMS 11

12 NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY Science Series 28

40a (38b). One pair of antennae; interstitial forms PROTODRILIDAE

1976 THE POLYCHAEtE WORMS 13

14 NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY Science Series 28

1976 THE POLYCHAETE WORMS 15

Ia. Two asetigerous anterior segments 2

16 NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY Science Series 28

lob (9a). Acicular spines absent in abdominal neuropodia Haploscoloplos

Generic Definitions include crenulated capillaries, hooks and intermediate

1976 THE POLYCHAETE WORMS 17

neurosetae include crenulated capillaries, hooks and Invalid Genera

18 NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY Science Series 28

5b (4b). Modified setae otherwise 6

Generic Definitions Nuchal organs prostomial; median antenna absent.

1976 THE POLYCHAETE WORMS 19

Generic Definitions (1971); it now is divided into two sub-families with

Questa Harman 1966a, Q. caudicirra Hartman 1966a;

FAMILY CTENODRILIDAE KENNEL 1882

Body short or long, prostomium without appendages,

20 NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY Science Series 28

2a (lb). Unpaired dorsal cirri present; branchiae absent Zeppelinia

Generic Definitions Setae include capillaries and short, thicker dentate or

Parergodrilus Reisinger 1925, P. heideri Reisinger

Stygocapitella Knollner 1934, S. subterranea Knollner

Prostomium without appendages, palps absent. At

1976 THE POLYCHAETE WORMS 21

Prostomium without appendages; a single peristomial

FAMILY COSSURIDAE DAY 1 963

Prostomium without appendages, a single median

22 NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY Science Series 28

Suborder Spioniformia FAMILY SPIONIDAE GRUBE 1850

1976 THE POLYCHAETE WORMS 23

24 NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY Science Series 28

1976 THE POLYCHAETE WORMS 25

Pygospio Claparede 1863, P. elegans Claperede 1863;

26 NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY Science Series 28