Professional Documents
Culture Documents
Science Series 28
February 3, 1977
ti
TABLE OF CONTENTS
PREFACE VII
ACKNOWLEDGMENTS IS
INTRODUCTION 1
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 FLABELLIGERIDA 1 15
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
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.
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;
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
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
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
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
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
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
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.
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
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
ORDER PSAMMODRILIDA
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
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
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.
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.
Key to Genera
1. Antenna frontal, first segment with series of long setae Poecilochaetus
2. Antenna median, first segment asetigerous Elicodasia
Invalid Genus
Suborder Chaetopteriformia
Key to Genera
Invalid Genera
Suborder Cirratuliformia
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
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
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
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.
Key to Genera
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
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-
Thorax with 12 segments; one asetigerous segment
lary setae in the notopodia and hooks in the neuropodia.
present; first setiger complete. Up to segment 6, cap-
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
Thorax with 12 segments; one asetigerous segment
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
present; first setiger complete. Up to segment 10, cap-
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
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
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
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
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
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.
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.
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
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
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 pros- tomium 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
Key to Genera
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
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
54 NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY Science Series 28
FIGURE 14. Family APHRODITIDAE, Aphrodita refulgida, Halfmoon Bay, California, felt removed on right side, 2.5x.
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.
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
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
86b (83b). At least some neurosetae bidentate 88
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
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
1976 THE POLYCHAETE WORMS 61
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; neuro- setae 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.
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
Key to Genera
FIGURE 17. Family SIGALIONIDAE, Sthenolepis japonica, Bay of Nha Trang, Viet Nam, 50 m, 25x.
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.
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
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
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
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
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
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.
Key to Genera
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
Brania Quatrefages 1866, Exogone pusilla Dujardin Eusyllis Malmgren 1867, E. blomstrandi Malmgren
1851; 21 species. 1867; 27 species.
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.
EUSYLLINAE. Three antennae; two pairs of ten-
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.
EUSYLLINAE. Three antennae; two pairs of ten-
Opisthodonta Langerhans 1879, O. morena Langerhans
tacular cirri. Occipital flap present. Pharynx with
1879; 2 species.
valve; teeth absent.
EUSYLLINAE. Three antennae; two pairs of ten-
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
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.
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
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
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
Key to Genera
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
,I
species.
Posterior segments biramous. Chevrons absent;
pharyngeal organs large and of several kinds. Noto-
setae knobbed or falcate hooded hooks; neurosetae
composite spinigers.
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
Invalid Genera
Eone Malmgren 1866, see Glycinde
Epicaste Kinberg 1866b, see Glycinde
Lacharis Kinberg 1866b, indeterminable
Leonnatus Kinberg 1866b, see Goniada
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
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.
Key to Genera
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
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
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
Invalid Genera
Key to Genera
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.
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
1976 THE POLYCHAETE WORMS 101
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
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
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
ORDER EUNICIDA
Superfamily Eunicea
Two to five pairs of lateral jaws (maxillae) and
usually one pair of lower jaws (mandibles).
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
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;
branchiae present. Setae include limbate setae, pectinate
setae, composite spinigers and subacicular hooks.
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
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
108 NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY Science Series 28
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.
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
Generic Definitions
Key to Genera
2
4
Drilognathus
3
Drilonereis
Notocirrus
Biborin
5
Arabella
6
Oligognathus
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-
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
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
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
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
Invalid Genera
ORDER FLABELLIGERIDA
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
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
Key to Genera
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
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.
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
44b (39b). Fourteen thoracic uncinigers 47
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
55b (53b). At least 12 thoracic uncinigers 58
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
58b (55b). Fourteen thoracic uncinigers 60
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
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.
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.
AMPHARETINAE. Three pairs of smooth branchiae;
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
Amphitrite O.F. Muller 1771, A. cirrata O.F. Muller Artacamella Hartman 1955, A. hancocki Hartman
1771; 17 species. 1955; 2 species.
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
1860; 3 species. 1860; 8 species.
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, re- present 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.
1976 THE POLYCHAETE WORMS 133
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
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
Key to Genera
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
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
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
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.
SERPULINAE. Seven thoracic setigers; operculum
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.
SERPULINAE. Seven thoracic setigers; operculum
Hydroides Gunnerus 1768, H. norvegica Gunnerus
spherical, opercular stalk smooth. Collar setae and
1768; 85 species.
side setae absent; all thoracic setae limbate, abdominal
SERPULINAE. Seven thoracic setigers; operculum
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.
SERPULINAE. Seven thoracic setigers; operculum
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
SERPULINAE. Seven thoracic setigers; operculum
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
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
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.
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).
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
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.
Key to Genera
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
1976 THE POLYCHAETE WORMS 153
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.
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
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
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).
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).
20
27
-50
-28
29 Pwl -52
mp
lo
11
Ficuan 41. Diagram of selected tens defined in the Glossary
#19-60.
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.
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).
1976 THE POLYCHAETE WORMS 161
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INDEX