The Polychaete Worms. Definitions and Keys To The Order, Families and Genera

THE POLYCHAETE WORMS
DEFINITIONS AND KEYS TO THE ORDERS, FAMILIES AND GENERA
THE POLYCHAETE WORMS Definitions and Keys to the Orders, Families and Genera
By Kristian Fauchald
NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY In Conjunction With THE ALLAN HANCOCK FOUNDATION UNIVERSITY OF SOUTHERN CALIFORNIA
Science Series 28 February 3, 1977
TABLE
PREFACE ACKNOWLEDGMENTS INTRODUCTION CHARACTERS USED TO DEFINE HIGHER TAXA CLASSIFICATION OF POLYCHAETES ORDERS OF POLYCHAETES KEY TO FAMILIES ORDER ORBINIIDA ORDER CTENODRILIDA ORDER PSAMMODRILIDA ORDER COSSURIDA ORDER SPIONIDA ORDER CAPITELLIDA ORDER OPHELIIDA ORDER PHYLLODOCIDA ORDER AMPHINOMIDA ORDER SPINTHERIDA ORDER EUNICIDA ORDER STERNASPIDA ORDER OWENIIDA ORDER FLABELLIGERIDA ORDER FAUVELIOPSIDA ORDER TEREBELLIDA ORDER SABELLIDA FIVE "ARCHIANNELIDAN" FAMILIES GLOSSARY LITERATURE CITED INDEX
OF
CONTENTS vii ix 1 2 7 9 9 14 19 20 21 21 31 41 45 100 103 104 114 114 115 117 118 135 152 156 161 180
Preface
THE STUDY of polychaetes used to be a leisurely occupation, practised calmly and slowly, and the presence of these worms hardly ever penetrated the consciousness of any but the small group of invertebrate zoologists and phylogenetlcists interested in annulated creatures. This is hardly the case any longer. Studies of marine benthos have demonstrated that these animals may be wholly dominant both in numbers of species and in numbers of specimens. In some areas they are even dominant in biomass. If any .statements are to be made about the biology of the benthic environments, some note must be taken of the polychaetes. Furthermore, the keeping of some of these animals in culture has proved feasible, and some polychaetes even have become famous for their value as test animals in polluted areas. The development of some polychaetes appears to be genetically interesting, and studies of evolutionary rates and genotypic and phenotypic adaptations in these worms may prove very illuminating to our understanding of the benthic environment. All of these developments make it important that the major morphological and anatomical features be well understood, and preferably that the terminology and taxonomic categories be agreed upon by most workers. This review is an attempt at summarizing current information about the taxonomy and morphology of these animals. During the reviewing process, it became painfully obvious to me how exceedingly poorly known the group is and how few generalizations can be made on the ecology and evolution of the polychaetes. I hope this review will bring forth a spate of investigations, by persons wishing to prove me wrong in my phylogenetic speculations, but also by persons willing to put in the hard work needed to fill in the gaps in our knowledge. I apologize to my fellow polychaete workers for introducing a complex superstructure in a group which so far has been remarkably innocent of such frills. A great number of very sound partial schemes have been suggested from time to time. These have been only partially considered. The discussion is complex enough without the inclusion of speculations as to how each author would have completed his or her scheme, provided that he or she had had the evidence and inclination to do so. Kristian Fauchald 19 May 1976 Los Angeles
Acknowledgments
THIS REvEW is a direct outcome of a lecture I gave during a class in polychaete biology at Santa Catalina Marine Biological Laboratory in 1973. It has benefited greatly from discussions with all of the students in the class, but perhaps especially Bill Kennedy was instrumental in pushing me into giving the lecture in the first place. 1 have also discussed parts of the paper with Ray Emerson, Tom Kauwiing, Fred Piltz and Bob Smith. My co-teacher at Catalina, Paul C. Schroeder, was a great help in formulating some of these ideas. Some of the ideas presented here were also discussed with the late Dr. Olga Hartman, whose sound advice was always valued by me. Dr. Pat Hutchings advised me of some errors in the terebellid key, for which I am grateful. The good illustrations were made by Ms. Catherine Link; the others I am responsible for. Ms. Jo Anne Woodcock expertly typed most of the manuscript, prepared the index and cleaned up my English where needed, for which I am grateful. I am also very grateful to Mr. R. Edward Ostermeyer for seeing the paper through. I would wish to thank Dr. Robert J. Lavenberg, Natural History Museum of Los Angeles County, for prodding me into writing this book in this format and for his constant encouragement and enthusiasm for what grew from a moderately long key to families to a rather more massive offering. I am also very grateful to Dr. Bernard C. Abbott, Allan Hancock Foundation, University of Southern California, for his support and enthusiasm and for letting me have the time to do the writing on this paper.

DEFINITIONS AND KEYS TO THE ORDERS, FAMILIES AND GENERA^
By
KRISTIAN FAUCHALD^
ABSTRACT; A review of the classification of the Class Polychaeta (Annelida) with comments on the characters used to identify the different included taxa has led lo the recognition of seventeen orders. All taxa down to the generic level are defined and a phylogenetic sequence suggested. Keys are presented to the families and genera of the Polychaetes.
INTRODUCTION Polychaeta is part of the old, diffuse concept Verines, a group that perhaps can best be defined as comprising all animals that are longer than wide and non-vertebrate. For the last seventy years or so, the polychaetes have been grouped with the oligochaetes and hirudineans and a few smaller groups into the phylum Annelida. This phylum contains .segmented, coelomate worms in which a secondary loss of segmentation may have taken place, but in which traces of such segmentation at least can be recognized internally. Other, more formal definition of the phylum can be found in standard text-books. Definition of the annelid classes has varied, however, everyday recognition of members of the three major classes has never been a problem: The Hirudinea contains the leeches, the Oligochaeta the earthworms and their allies, and the Polychaeta marine worms, such as the sand worm (nereids) and bloodworms (glycerids) used for bait in parts of the world. The three classes appeared also, at least grossly, to be separated ecologically in that the leeches were supposed to be parasitic, the earthworms terrestrial and the polychaetes were most common in the marine environment. This separation is obviously unsatisfactory and as information about these animals accumulated towards the end of the last century, most workers settled on a grouping which associated the leeches and the earthworms with each other in one group opposed to the polychaetes. Members of both the former classes are hermaphroditic and have various complex glands associated with reproduction. Furthermore, both leeches and earthworms lack parapodia (fleshy unjointed segmental appendages)
and the setae, if present, only rarely occur in bundles. These two groups were considered more advanced than the marine, dioecious polychaetes. The polychaetes have been defined for the last seventy years as dioecious, marine annelids with parapodia bearing numerous setae. They also should have anterior appendages of various sorts (antennae, palps, tentacular cirri) and the gonadal ducts should be simple. These definitions work if some of the smaller groups are disregarded. If these groups are taken into account, as they must, the only separation that consistently can be made between the oligochaetes/leeches and the polychaetes, is the presence in the former grouping of hermaphroditic gonads limited to a few segments. Some hermaphroditic polychaetes are known, but these usually have gonads in a large fraction of the total number of segments. It is then difficult to give a good, consistent and practically useful definition of what is meant by a polychaete, but a definition along the lines suggested below, should separate them from the other annelids with reasonable accuracy. The polychaetes are multi-segmented annelids with parapodia; setae are present in distinct fascicles. They are dioecious and have simple exit ducts from the
'REVIEW COMMITTEE FOR THIS SCIENCE SERIES

ROBERT J. LAVENBERG
DONALD REISCH
Âllan Hancock Foundation, University of Southern California. Los Angeles, California, 90007. Contribution Number 358 of the Allan Hancock Foundation.
NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY gonads. They are usually marine, more rarely freshwater and only rarely terrestrial or parasitic in habitat. Any of these features need not be present and none of them is essential for the recognition of an animal as a polychaete. This topic has been treated in considerably greater detail by Clark (1969) and to a lesser extent by Fauchald (1974a). A key morphological feature and at the same time one of the most important taxonomic characters of the polychaetes is the setal (chaetal) construction. The setae are ectodermal derivatives, formed by ectodermal cells that during the development have migrated to a position well below the rest of the ectodermally derived epidermis. Each seta consists of a bundle of filaments laid down by a basal chaetoblast and up to several lateral cells. The material in the setae is a glycoprotein, consisting of chitin (a polysaccharide) and a protein cross-linked at the time of formation. The formation of structural details in the setae is very well controlled, but exactly how this takes place has only partially been clarified. The best current review of this topic was 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 Milne Edwards 1834, pp. 24-26). The separation is based on the development of the anterior end and the life habits of the included species. The errants are supposed to have a large number of equal body-segments. The anterior appendages are few in number and differentiated into palps, antennae, tentacular cirri, etc. These worms are considered freeliving and, generally, should be rapacious in habits. All polychaetes with jaws are included in this order; thus the onuphids, despite their tubicolous habits, are considered errants since their large jaw-apparatus resembles the jaw-apparatus in other, non-tubicolous eunicidlike animals. The sedentaries are supposed to have a limited number of body segments. The body may be separated into different regions. Anterior appendages may be absent or a few to many similar appendages may be present. The sedentaries have short parapodia associated with their tubicolous or burrowing habits and are usually deposit- or filter-feeders. These definitions have not changed much over time (cf. Grube 1850, p. 281 and tables; Fauvel 1923a, pp. 27-29; Hartmann-Schrder 1971, p. 29). The advantage of the system is that the bulk of the 8,000-1described species of polychaetes separates into two roughly similar groups in terms of numbers of species and genera as well as families. The separation is otherwise unsatisfactory since neither order can clearly be defined. Several attempts have been made to subdivide the polychaetes in a more acceptable manner (Dales
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1962, pp. 424-425; Clark 1969, p. 47). Polychaete taxonomists have tended to disregard these attempts and have continued to treat the polychaetes as if the class consisted of two orders (Fauvel 1958, pp. 166190; Hartmann-Schrder 1971, p. 29) or subclasses (Uschakov 1955a), or have treated the group as if it consisted of about 75 distinct and unrelated families (Hartman 1968, 1969). The problem with all proposed schemes is that they are internally inconsistent. Furthermore, they give no better solutions to classificatory problems than the old, admittedly anificial, separation into two orders. The three most ambitious recent proposais were by Dales (1962), Storch (1968) and Clark (1969). Dales used the variable structures of the eversible stomodeal region (pharynx) to separate different groups. The arrangement of the body-wall musculature was used by Storch. Clark used a variety of different structures to characterize his eight orders. These authors gave no formal definition of any taxon above the family level (except by inference from contained taxa) and it has been difficult to evaluate their schemes.
CHARACTERS USED TO DEFINE HIGHER TAXA Major anatomical and morphological features were reviewed during a study of the phylogenesis of the polychaetes (Fauchald 1974a). Below is given a survey of the findings with an expanded discussion of their taxonomic aspects. A. Prostomium. The prostomium usually is distinct and may have or lack appendages. In several families it is more or less fused with the peristomium and the first segments. The degree of fusion is difficult to determine even in an examination of the nervous system so the degree of distinctness of the prostomium is a character that can have no great taxonomic value (see Benham 1894, 1896). Prostomial appendages include antennae and palps. Antennae are innervated through single roots directly from the brain; palps always have double roots, either from the brain or from the circumesophageal ring (kesson 1963; Orrhage 1966). Antennae are always sensory; palps may be sensory or may be used as feeding appendages. The presence of either one or both categories of appendages is considered here of great importance. The position of the palps varies from ventral to dorsal, from frontal to occipital, The position and function of the palps furnish important taxonomic characters. It is impossible to distinguish any other classes of prostomial appendages either on morphological or anatomical grounds. B. Peristomium. The larval peristomium is the immediate prototrochal region of the trochophora larva;
1976
it may persist as the adult peristomium at least in some forms and it appears to be completely pre-segmental In nature, at least in some forms (Altesson 1967). However, the structure called the peristomium in most taxonomic studies consists of a fusion of this larvally derived structure and one or more true segments. The larvally derived peristomium may carry a single pair of dorsal cirri called peristomial cirri. The fused segments may carry parapodial remnants called tentacular cirri. The number of tentacular cirri vary from one to four pairs in the hesionids; other families tend to have
PALP ANTENNA
PROSTOMIUM PERISTOMIAL CIRRUS PERISTOMIUM SEGMENT liMticaroHil SETIQER PARAPODIUM SETA SEG MENTAL
POSTSEQMENTAL
PYGIDIUM ANUS ANAL CIRRUS
NOTOPODIUM
EUROPODIUM VENTRAL NERVE PARAPODIUM
I. Diagrams showing the major morphological features of a generalized polychaete.

FIGURE
a constant number, or at least only a few alternative numbers (nereids, syllids, phyllodocids, etc.). Fused segments are often present, even if tentacular cirri are absent. It would be valuable to distinguish the two kinds of peristomia, but current usage seems ingrained and little would be gained by coining a new term; it should, however, be remembered that the current term may cover two very different structures. C. Eversible pharynx. Most polychaetes can everse 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 (Eunicea, Amphinomida, etc.); others have an axial pharynx that is developed symmetrically, or at least nearly symmetrically. The axial pharynx may be followed by a strongly muscular region (nereids, nephtyids, glycerids, etc.) or this musculature may be absent (arenicolids, maldanids, etc.). Usually each family has a characteristic kind of pharynx, but in some families (e.g., Spionidae) both plate-muscle and axial pharyngs are present (Orrhage 1966). Some plate-muscle pharyngs are poorly muscularized and may be difficult to distinguish from weakly developed axial pharyngs so the apparent overlap in distribution of the two kinds of pharyngs 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 precisely to other variable morphological features and to environmental variables (Fauchald and Smith, in preparation). D. Parapodia. Polychaete parapodia can be biramous, with both note- 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 various parapodial lobes and cirri is very important at the generic and specific levels. The presence of branchiae associated with the parapodia is of variable importance. The presence of branchiae may not even be considered 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 Branchioasychis). Generally, however, the presente of branchiae is a specific character, but with generic
NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY
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importance where warranted by evidence. Branchiae are of sufficient biological importance, so fairly good evidence must be presented to demonstrate that these structures are of less than specific importance. E. Setae. Numerous kinds of setae have been described (uncini, limbate setae, pectinate setae, subacicular hooks, composite spinigers and falcigers, etc.). The basic kind of setae in each parapodial ramus is usually a family character, but not uniquely so. Thus, all phyllodocids have composite setae, but not all polychaetes with composite setae are phyllodocids. The detailed construction of the setae is important at the specific level. The importance of accurate examination of the setae still is underestimated by most taxonomists; precise observations require close microscopic work to elucidate them and this kind of work may be necessary even in routine identifications. Moreover, recent studies with scanning electron microscope have demonstrated clearly the importance of accurate work on the setae (Thomassin and Rcard 1972). F. Nephridia. The structure and distribution of nephridia have been used at the subfamilial and generic levels (Hessie 1917 on terebellids). The character may be of wider usefulness at higher levels, but the variation in these features has been too little studied in most families to make the character useful at the present time. Most polychaete families are characterized by a complex set of features and cannot be identified by reference to a single structure. The proposed taxonomic schemes have failed because they did not take this into account. They all were based on two or perhaps three manifestations in one important structure. By defining each manifestation precisely, apparent "intermediate" forms appeared and had to be included in one or another category as an exception or be left outside the proposed system. Usually, the "intermediate" form would be loosely appended to one taxon or another; the definition of the taxon would then be left intact. It became impossible to find these "hidden" taxa in the system.
SOME USEFUL TECHNIQUES
The following comments are necessarily incomplete and are meant only as a first-hand guide to work on polychaetes, especially the handling necessary to perform identificatory work, excluding the handling necessary for other kinds of investigations on either live or dead polychaetes. The main topics covered are collection, screening, fixation and preservation and the most common techniques in laboratory handling. Collection.Polychaetes are found nearly everywhere in the marine environment and thus can be caught with every kind of gear imaginable. They are soft-bodied animals, and hand-collected specimens
tend to be rather poorly treated. The most complete, and thus most easily identifiable specimens will be caught with gear that takes chunks of the environment and in situations where the animals are allowed to crawl out, or where the material is gently screened. Even in S.C.U.B.A. collection it is advisable to collect masses of material, rocks, seaweed tufts, sandsamples, etc. and put them in separate plastic bags or similar waterproof sample bags for later treatment. Most polychaetes are small and active animals; it is therefore useless to put a bottom sample in the ordinary game-bags used in diving, since most of the worms will escape before the sample can be treated further. Shipboard sampling can be done with any of several kinds of a series of gear, quantitative as well as qualitative, If dredges are used, I recommend hauls as short as practicable, since the churning of the material in the dredge will tend to grind up the polychaetes or disturb them enough to make them autotomize appendages and, often, the whole posterior end. Samples of hard substrates, algal mats, rubble, etc. should be put in a large container of some sort, covered completely with sea water and be left standing undisturbed for several hours. The samples should be placed in the dark or at least in the shade and should be kept cool, though not necessarily refrigerated below the ambient temperature at the samplmg site. It is especially important to leave the sample in the dark if it contains algae in quantity. As the oxygen concentration in the water decreases, the polychaetes will leave the substrate and congregate around the rim of the vessel at the air-water interface. They can easily be scooped from the surface with either a small screen (0.5 mm mesh-size) or simply a spoon. If the sample contains a large number of motile, large animals, such as crabs or brittle-stars, it is best to remove most or all of these as soon as possible. The samples should be left standing for several hours (212 hours) depending on the lighting, the quantity of water in relation to sample size, the temperature, etc. After the samples have been treated in this manner, the substrate should be carefully sorted throughplucked to pieces if necessaryand the water screened for animals that left the substrate but did not reach the surface. This whole process should be done as quickly as possible, since polychaetes deteriorate very rapidly after death. The method will not quantitatively remove polychaetes, since some polychaetes are unable to leave their tubes or burrows and others generally do not approach the surface, even when the water becomes very foul. The process can be speeded up by adding 7% MgCla, but we have not found this to be any great advantage. Screening.Samples of soft substrates, such as sands and muds, must be screened. Quantitative benthic studies now routinely use I mm screens, but these undersample the polychaetes badly, both in numbers of
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species and in specimens. The use of 0.75 mm screens does not improve matters greatly, but 0.5 mm screens appear to catch most polychaetes quantitatively. In Inshore areas, the sands and gravel will make the use of 0.5 mm screens rather impractical, but if a complete survey of the fauna is contemplated, this cannot be avoided. Thus, the objective of the study undertaken must be considered carefully. I would generally recommend that, for each area studied, at least some samples be screened with a double set of screens, both I.O and 0.5 mm, and the results compared so that the level of inaccuracy engendered by the use of 1.0 mm screens can be estimated. This has to be done for every major sediment type and cannot be guessed at from one sediment type to the next, nor from one geographical location to the next. This is because the fraction of small species appears to vary geographically. Samples should under any circumstances be washed gently with very large quantities of water. It is useful to have the water prescreened so that pelagic organisms can be avoided, and deep-water samples should be treated with water as cold as possible. Nothing is gained by hurrying the screening process, but each sample should be screened as soon as possible after getting it on deck. Deep-water samples that cannot be screened immediately should be refrigerated. It is of the utmost importance for good results that the screening process be done carefully; poorly screened samples contain a large number of mangled specimens, and such specimens usually cannot be identified. The net result will be a waste of sampling time and effort and, not least, loss of time needed for identification of the specimens after the samples have been returned to the laboratory. In terms of the time and effort needed for the different parts of the processing of a single sample, we now generally calculate that on the average it takes about 50 minutes of shipboard time to take and process a single shelf sample with a box-core and about three weeks of manhours to adequately treat and interpret this sample in the lab. Thus, the few minutes of ship time to be gained in preparing the samples poorly will be offset by the need for a larger number of replicate samples to get adequate numbers for mathematical treatments in the lab. Once the sediment has been removed from the screens as completely as possible, the retained material should be chased down to one side of the screen with the help of a gentle stream of water applied to the outside of the screen. Do not sort or pick through the material on the screens: non-preserved polychaetes should be handled as little as possible and never moved at all except with the help of a gentle stream of water applied to the outside of screens or elsewhere. Fixation and Preservation.For the last few years we have routinely narcotized the whole sample as retained on the screens. The material on the screens is washed down onto a large enamel pan with as little sea
water as possible. The pan is then flooded with a solution of 7% MgCU in sea water and allowed to stand for about half an hour. Then the contents of the pan are poured carefully through a screen and transferred, with as little water as possible, to a sample jar. The narcotizing solution can be used several times if so desired. Standard fixating agent for polychaetes is 10% neutralized formalin in sea water. The most commonly used neutralizing agent is borax (Na2B407, technical grade). The sample should fill no more than one-third of the sample jar and the jar should be completely filled with the formalin solution, The jar should be capped and gently but thoroughly inverted several times to get complete mixing. Allow the sample to settle, decant off about one-half of the solution and fill the jar with fresh formalin solution. It is much better to split a sample into several jars than to fill one jar completely with the entire sample and get incomplete and imsatisfactory fixation. Histological fixatives can also be used on bulk samples, and in general such fixation is better on small, fragile polychaetes if done prior to any sorting. If such fixatives are used, the ratio between sample and fixative must be even lower than the one indicated above, and the fixative should be changed twice in rapid succession to avoid dilution effects. Samples should be left in formalin for at least 24 hours and can be left in the fixative for several weeks. However, after 24 hours, the samples are ready to be transferred to the preservative, usually 70% isopropyl or ethyl alcohol in distilled water. Before transfer, the samples must be washed in fresh water once or twice to remove the salt; if this is not done, setae and other details will become the crystallization sites for salt crystals, and these are difficult to remove after they have formed. One change of preservative is necessary to ensure full strength. DO NOT ATTEMPT TO USE ETHYL ALCOHOL AS A FIXATIVE, even if sensitivity to formalin becomes a problem. The specimens become completely unusable after a short period of time, and again the fraction of unidentifiable specimens goes up drastically. Postfixation with formalin of material originally fixed in alcohol does not work. Specimens treated as recommended above usually retain most of their appendages, and a large fraction of complete specimens is usually present. However, a certain number of incomplete specimens are to be expected in any treatment, especially with standard narcotizing time as suggested above. The treatment suggested is rather more elaborate than usual, but we have found that we aie amply rewarded by a much higher than usual fraction of identifiable specimens. Thus, less replication of samples is needed in benthic surveys and this in itself represents a savings in both time and money.
NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY Laboratory Treatment of Samples, Identificatory Techniques.The equipment needed to identify polychaetes includes one stereo microscope per person and one compound microscope per two persons, as the minimum. The compound microscope must be capable of magnifications up to 1100 times, and the stereo microscopes should have magnifications to about 100 times. A focusabe microscope lamp is necessary for use with the stereo microscope. Each person will also need two pairs of watchmaker's forceps, two needles (we use insect pins glued to applicator sticks with epoxy glue), fine scissors, a small scalpel (we use pieces of razor blades glued to applicator sticks with epoxy glue), a small bottle of glycerol alcohol mixed with one-half glycerol and one-half 70% alcohol, a bottle of immersion oil, depression slides and flat slides, cover slips, petri dishes of various sizes (preferably with tight-fitting lids), cotton, paper, and #2 pencils (HB or F works well). The sorting and identification of polychaetes is a two-step operation. Each sample should first be sorted to family under a stereo microscope. The samples must be sorted while completely submerged in alcohol, and since light from the microscope lamp is apt to evaporate the alcohol it should be refi^shed iiom time to time. Polychaetes always have to be treated while completely submerged, and after being transferred to alcohol cannot be left dry for more than a few minutes at a time. Small polychaetes tend to dry out very quickly, and dried out polychaetes are largely unidentifiable. Sorting can be done directly into vials completely filled with alcohol, and each vial receives a label with the name of the contained taxon, as well as with the station number for that particular sample. It is imperative to do this immediately rather than having to remember the content and position of each vial later. Use good, high-rag-content paper of sufficient weight (20 pound or higher) and a good pencU. The labels should be big enough to stand up in the vial unsupported, but not so large as to cover the contents, and they should end up well below the upper margin of the vial. We prefer to use straight-sided vials, capped with cotton plugs, and to store these in jars filled with alcohol. Screw-capped vials almost invariably have a shoulder which makes it difficult to remove specimens and labels when needed. Screwcaps normally are not air tight, so the vials will have to be stored within a larger jar regardless. After the sample has been sorted to family, we collect the members of each family in a single jar. Thus, the sample is no longer intact as such, but has been distributed taxonomically among families. It is thus of very great importance that careful and accurate notes be taken on the numbers of vials for each station, so that later the station can be reconstructed accurately on paper. This is done because we have found it con-
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siderably more efficient to identify the polychaetes family by family in larger numbers. It is much easier to compare specimens from different samples, and the number of dubious identifications can be decreased drastically by this means. If the samples are to be stored as units, the vials can always be reunited afterwards if desired. When polychaetes are being identified, certain standard observations should always be made. Always find the anterior end and take note of the number and arrangement of anterior appendages. Scan the body for obvious differences in parapodial structures and for such features as the position and number of branchiae. Most specimens preserved as indicated above will be relatively easy to handle with two pairs of forceps, and it is usually much easier to move the specimen than to move the dish. This sort of scan is usually all that is needed to get a polychaete to family. To identify the animals to genus and species, more accurate observations are usually necessary and various dissections must be performed. It is always necessary to remove a parapodium, if nothing else, because a good look at the setae is needed. This can be done with the use of two pairs of forceps, a scalpel or iris scissors. For most medium-sized polychaetes in good condition, just pulling off a parapodium with a pair of forceps is the easiest method. Care must be taken that both parapodial rami and associated cirri and branchiae come off. Some workers find it easier to use scissors or scalpel, and these instruments must be used on poorly preserved specimens or on larger animals. The parapodium should be mounted on a slide; larger ones must be mounted on depression slides, but normally a flat slide is better, since depression slides cannot be used with high-power compound microscopes. Generally, the parapodium should be mounted with the anterior side facing the observer; however, in certain families and genera, a posterior view may be more informative. Before mounting the parapodium, look at some appropriate parapodial illustrations showing the features to be observed. A parapodium mounted in a dorso-ventral position gives no more information than does looking at the whole animal, so a mount showing both notopodium and neuropodium is necessary. We use glycerol-alcohol for these mounts and only rarely make permanent mounts. Which parapodiimi should be removed depends on which family is being studied. Generally, a median parapodium from a long series of similar-looking parapodia will be best, but in special cases the anteriormost or one specific parapodium will have to be removed to study some specific detail of importance in that taxon. Thus, in members of the genus Pista, for example, anterior, median and posterior thoracic parapodia must be removed, and in the genus Magelona
1976
the ninth parapodium must be detailed. Again, it is wonhwhile to look through the key to be used before deciding which parapodium and, in some cases, how many parapodia, should be removed. It is also worthwhile to scan the body of the animal carefully before deciding on a specific parapodium, to see that the setae are as complete as possible so no extra mounts of setae become necessary. Setal distribution varies from one group of polychaetes to another; if it is possible to remove one single parapodium and get all setal types represented at once, this saves wear and tear on the specimens and also saves time in preparation. Generally, parapodial structures are most easily observed at relatively low magnifications under the compound microscope, but critical decisions, especially on structures on top of a thick preparation, may be most easily followed at higher magnifications. Setal structures, especially the presence of fine hairs along the cutting edge of simple setae, are best seen under oil immersion with lOOx objectives. It does not help 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 immersion oil also between the condensor and the lower side of the slide. Be sure to adjust the lighting of the compound microscope every time the magnification is switched; it gives a much more satisfactory picture of the structure and will also, in the long run, save the eyesight of the worker. Dissection of the anterior end of the polychaete may also be necessary for observation of the structure of the eversible pharynx or jaw structures. Important structures are situated in the midline, dorsally or ventrally or both; thus, a dissection should avoid cuts into the midlines of the animals. We generally make a longitudinal slit on the dorsal side, well lateral to the midline. The length and position of the slit will depend on the purpose of the dissection and on the relative position of the eversible pharynx. The pharynx may be preserved in the completely retracted position or in various stages of eversin; jaw structures are usually situated anterior in the body, even in forms with a long pharynx. If the purpose of the dissection is observation of the lining of the pharynx in one of the forms with a long pharynx, the slit can be made farther back than usual. We usually continue by making transverse slits at both ends of the longitudinal one. This makes it possible to lift a flap of tissue containing the body-wall proper. We have found it most useful to leave the pharynx and the jaws in situ as much as possible; they are almost invariably lost if removed. Especially in the case of the complex jaw structures of the eunicean polychaetes, it is important to treat them in a similar manner in all specimens, so that they are all flattened to a similar degree when the observations are being
made, etc. The jaws in these forms are always observed from the dorsal side and the jaw formulae are given from the posterior to the anterior end, the left jaw being mentioned first in each formula. The pharyngeal lining has characteristic structures in members of several families; it is usually not necessary to remove the lining in nephtyids and phyllodocids, but in order to characterize the lining of members of glycerids and goniadids, this must be done. It is of great importance that the lining be well oriented in the goniadids, since the position of the different kinds of pharyngeal organs is considered of taxonomic importance. The fine structure of the glycerid pharyngeal organs cannot be seen except in oil immersion, and critical lighting is of the utmost importance for a clarification of these structures. Parts that have been dissected out are best put in a small, separate vial stored within the larger vial with the specimens. Leaving small parts loose in the vial will invariably lead to their loss.
CLASSIFICATION OF POLYCHAETES The scheme proposed below (cfr. also Table 1), is based on phylogenetic ideas presented elsewhere (Fauchald 1974a). The sequence of families indicates an increasing morphological distance from the ancestral polychaete as this was defined on that occasion, but since the several orders and families are considered the results of a rapid radiation in Pre-Cambrian to Cambrian times (Fauchald 1974a), this sequential arrangement can only poorly represent the phylogenetic pattern. Major anatomical as well as morphological features were used to define the orders. This may make them difficult to use in practical taxonomic work. The framework formed by including the anatomical features is more satisfactory in that each order now can be defined to exclude all non-members. Theoretically different evidence should be used to define each taxonomic level. One feature (e.g., the structure of the eversible pharynx) once used at one taxonomic level, should not be used at another (lower) level within the same classificatory plan. This sort of separation was attempted here, but was only partially successful, in that one feature (e.g., the structure of the eversible pharynx) may have been used at the family level in one order, but at the sub-ordinal or superfamilial levels in other orders. Suborders have been recognized only where warranted; no attempts were made to create intermediate categories in all orders. Families not included in any intermediate category are listed alphabetically at the end of each order. The sequence of families otherwise indicates phylogenetic relationships within each order.
NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY 1 Survey of Polychaete Orders, Suborders, Superfamilies and Families.
TABLE
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1. O. ORBINUDA F. Orbiniidae F. Paraontdae F. Questidae 2. O. CTENODRILIDA F. Ctenodrilidae F. Parergodrilidae 3. O. PSAMMODRILIDA F. Psammodrilidae 4. O. COSSURIDA F. Cossuridae 5. O. SPIONIDA SO. Spioniformia F, Apistobranchidae F. Spionidae F. Magetonidae F. Trochochaetidae F. Poecilochaetidae F. Heterospionidae SO. Chaetopteriformia F. Chaetopteridae SO. Cirratuliformia F. Cirratulidae F. Acrociiridae 6. O. CAPITELLIDA F. Capitellidae F. Arenicolidae F. Maldanidae
O. OPHELUDA SO. Glyceriformia F. Opheliidae F. Glyceridae F. Scalibregmidae F. Goniadidae O. PHYLLODOCIDA F, Lacydoniidae SO. Phyllodociformia Suborder not F. Phyllodocidae recognized: F. Alciopidae F. lospilidae F. Lopadorhynchidae F. Nephtyidae F. Pontodoridae F. Sphaerodoridae SO. Aphroditiformia F. Tomopteridae SF. Aphroditacea F. Typhloscolecidae F. Aphroditidae 9. O. AMPHINOMIDA F. Poiynoidae F. Amphinomidae F. Polyodontidae F. Euphrosinidae F. Pholoididae 10. O. SPINTHERIDA F. Eulepethidae F. Spimheridae F. Sigalionidae no. EUNICIDA SF. Chrysopetalacea SF. Eunicea F. Chrysopetalidae F. Onuphidae F. Palmyridae F. Eunicidae SF. Ksionacea F. Lumbrineridae F. Pisionidae F. Iphitimidae SO. Nereidiformia F. Arabellidae F. Hesionidae F. Lysaretidae F. Pilargiidae F. Dorvilleidae F. Syllidae Super-family not F. Calamyzidae recogaized: F. Nereidae F. Histriobdellidae F. Antonbruunidae F. Ichthytomidae
12. O. STERNASPIDA F. Slemaspidae 13. O. OWENIIDA F. Oweniidae 14. O. FLABELLIGEREDA F. Flabelligeridae F. Poeobiidae 15. O. FAUVELIOPGIDA F. Fauveliopsidae 16. O. TEREBELLIDA F. Sabellariidae F. Pectinariidae F. Annpharelidae F. Terebellidae F. Trichobranchidae F. Bogueidae 17. O. SABELUDA F. Sabellidae F. Sabellongidae F. Caobangidae F. Serpulidae F. Spiiorbidae Families of uncertain afniies: F. Dinophilidae F. Nerillidae F. Polygordiidae F. Protodriidae F. Saccocirridae
All known taxa to the generic level have been defined. Some taxa can be characterized by a single unique structure. The absence of this structure in all other taxa at that level has been left unstated to save space. Preferably, keys should have been made first to order and within each order, to famUy. For several reasons, this approach was abandoned. Identification of orders may be possible only after a detailed anatomical study; the orders are justifiable scientifically, but rather difficult in practical taxonomic work, as mentioned above. The general approach to keys taken here is that they are tools to make it possible to identify the taxa swiftly and easily and with as little damage to the specimens as possible. For that reason one master key to families is proposed. Furthermore, all keys were made reversible. If one has a fairly good idea of what kind of animal one has at hand from an illustration or otherwise, one can work backwards into the key to check the identification. The keys were made strictly dichotomous; trick wording was avoided. However, as in all other keys, the usage of adjectives and adverbs is rather different from common English usage. The family key was intended to discriminate between specimens that had lost deciduous features such as branchiae or antennae, etc., in addition to complete specimens.
Because of this and because of the general variability of key features within each family, several families key out at different points. The generic keys are dependent on the presence of deciduous features. Most of these keys are short enough to allow scanning of the total key for necessary corrections if such features have been lost. Under all circumstances, identifications made through the key should be checked by using the definitions of genera given. Introduction of a genus into the faunal lists should not be attempted based on identifications made through the keys in this paper. For publication purposes, the original literature must be consulted. A name erroneously introduced in the literature for any area is in practice indelible. Sloppiness has been the cause of more errors in the polychaete literature than all other causes combined. The keys and definitions given are wholly inadequate as a base for description of new genera, especially in the larger families. The keys are intended as aids in getting a first approximation in identification. Once achieved, a series of very valuable regional handbooks are available. Such include Uschakov (1955a and 1972), Hartmann-Schrder 1971), Hartman (1968, 1969), Imajima and Hartman (1964), Fauvei (1953), Day (1967) and Banse and Hobson (1974). Older, but
1976
still indispensable are the handbooks by Fauvel (1923a, 1927), Friedrich (1938) and Berkeley and Berkeley (1948, 1952). These books can be used for a second approximation to the identifieatory problems. However, before publication, the original research papers must be consulted. The review below is separated into several distinct parts. The family key is followed by an order by order review of all the families. For each family, a definition may be followed by a brief note, especially noting features useful in field identification. Major recent reviews are also mentioned at this point. Then comes a key to genera and definitions of all contained genera; followed by taxonomic notes. These explain new taxonomic combinations and taxa and specific positions taken in this paper. The type-species is named for all genera and an approximate number of species is given. Finally, a list of invalid genera has been added. Illustrations are given for one member of each family; as much as possible, identification features have been illustrated. A glossary contains most of the terms used, except those in general usage in invertebrate zoology; where necessary, small line drawings have been added to the glossary to illustrate idiomatic usage, etc. The literature cited contains references to the original descriptions of all genera listed, except the invalid ones and to major revisions and handbooks. It should be noted that this paper contains little that is wholly new; in most cases I have followed the most recent major revision of any family, or followed clues indicated by revisions currently under way. It is hoped that this compilation of information may make it simpler to get more complete revisions made for each family where needed. However, such a revision must be based on materials, not on descriptions. A major revision is a long and very laborious process, but must be undertaken; it is wholly unsatisfactory to base such revisions on previous descriptions only, since interpretation of descriptions frequently is dependent on poorly understood and used terminology; a fact that frequently has obscured close similarities in structure (cfr. Fauchald and Belman 1972; Blake 1975). Ultimately, one would hope that by organizing and defining each known taxon as clearly as possible, it
will be easier to debate the classification and phylogeny of pclychaetes sensibly; as stated by Clark (1969), the polychaetes do present an intractable problem of phylogeny for the time being. ORDERS OF POLYCHAETES The following polychaete orders are recognized: ORBINHDA, CTENODRILIDA, PSAMMODRILIDA, COSSURIDA, SPIONIDA, CAPITELLIDA, OPHELIIDA, PHYLLOE)OCIDA, AMPHINOMIDA, SPINTHERIDA, EUNICIDA, STERNASPIDA, OWENIIDA, FLABELLIGERIDA, FAUVELIOPGIDA, TEREBELLIDA, SABELLIDA. Members of the old order ERRANTIA are separated into three orders, PHYLLODOCIDA, by far the largest with most of the well-known families, AMPHINOMIDA and EUNICIDA. In addition, the small ectoparasitic spintherids have been assigned to their own order. The bulk of the recognized orders thus comes from the old collective group (or order) SEDENTARIA. This group was never adequately defined, because two very disparate subgroups had to be included; the highly modified species now included in the orders STERNASPIDA, OWENIIDA, FLABELLIGERIDA, TEREBELLIDA and SABELLIDA, and the simple-bodied forms now included in the orders ORBINIIDA, CTENODRILIDA, PSAMMODRILIDA, COSSURIDA, SPIONIDA, CAPITELLIDA and OPHELIDA. The latter seven orders contain structurally rather simple forms, but this should not be taken to indicate that the orders for that reason are related closely to each other. These forms are about as far apart as any other grouping of polychaetes that might be proposed, judging from differences in tagmatization, parapodial development and setal distribution. They could have been included under the old concept DRILOMORPHA (cfr. Uschakov 1955a, Dales 1962; Clark 1969), but defining this concept would have been very nearly impossible. The approach taken here, was that major different body constructions were given the rank of order and that modifications on these major body plans were given familial rank. Intermediate taxa were employed only where appropriate.
KEY TO FAMILIES
la. lb. 2a (la). 2b (la). 3a (2a). 3b (2a). 4a (lb).
External segmentation and setae absent 2 External segmentation and/or setae present 4 Paired antennae on the prostomium 3 Paired antennae absent; tactile hairs along the body; small interstitial forms DINOPHILIDAE Body long and slender; interstitial forms PROTODRILIDAE (in part) Body short and saclike, pelagic POEOBIIDAE Body a flattened disc with indistinct segmentation; ectoparasitic SPINTHERIDAE
10 4b (lb). 5a (4b). 5b (4b). 6a (5a). 6b (5a). 7a (6b). 7b (6b). 8a 8b 9a 9b (7b). (7b). (8a). (8a).
Science Series 28
10a (8b). 10b (8b). 11a 1 lb 12a 12b (5b). (5b). (11a). (1 la).
13a (lib). 13b ( 1 lb). 14a (13b). 14b (13b). 15a (14b). 15b (14b), 16a (15a). 16b (15a). 17a (16b). 17b (16b). 18a (17b). 18b (17b). 19a (18b). 19b (18b). 20a (15b). 20b (15b). 21a (20a). 21b (20a). 22a (21a).
Body not a flattened disc; segmentation usually distinct, if indistinct, then body clearly longer than wide 5 Dorsura 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 Dorsum without elytrae, elytral scars or felt 11 Neuracicula distally hammer-headed EULEPETHIDAE Neuracicula distally pointed 7 Prostomium with a single median antenna; dorsum with felt, or notosetae harpoon-shaped or held erect over the dorsum APHRODITIDAE Prostomium with one to three antennae; dorsum without felt; notosetae usually distinctly lateral in position, never harpoon-shaped 8 Neurosetae composite 9 Neurosetae simple 10 All posterior segments with elytrae; prostomium with one to three antennae SIGALIONIDAE Elytrae alternate with dorsal cirri along the whole length of the body; one antenna present PHOLOIDIDAE Spinning glands present; median antenna, if present, attached near the posterior or middle of the prostomium; notosetae absent POLYODONTIDAE Spinning glands absent; median antenna attached at the anterior margin of the prostomium; notosetae usually present POLYNOIDAE Notopodia with expanded, golden or brassy setae that more or less cover the dorsum 12 Notosetae otherwise (may be absent) 13 Prostomium with large facial tubercle and a median antenna; notosetae in rosettes .. PALMYRIDAE Prostomium without a facial tubercle; paired lateral and a median antenna present; notosetae in transverse rows CHRYSOPETALIDAE Posterior end covered ventrally by a chitinized shield STERNASPIDAE Posterior end not covered by a shield 14 Prostomium completely retracted between the first parapodia which have three pairs of tentacular cirri, partially supported by acicula PISIONIDAE (in part) Prostomium not completely retracted between the first parapodia which are otherwise equipped .. 15 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 Anterior end without exceptionally long, specialized setae (NOTE: Short, strong hooks may be present) 20 Specialized setae long and chambered, forming a protective cage around the retractable anterior end; body with numerous epithelial papillae FLABELLIGERIDAE (part) Specialized anterior setae do not form a protective cage; anterior end not retractable; skin-papillae few and small, or absent; setae otherwise 17 Specialized setae slender, distally curved, often spinous; prostomium with seven antennae ONUPHIDAE (part) Specialized setae stout, smooth and not distally curved; prostomium without appendages or with numerous tentacles 18 Specialized setae in a transverse row; tube conical, usually formed of closely fitted sandgrains PECTINARIIDAE 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 Specialized setae form paleae; anterior end with two to four pairs of branchiae . AMPHARETIDAE (part) Specialized setae form an operculum; anterior end without branchiae SABELLARIIDAE Anterior end, including in part the prostomium, transformed into a tentacular crown 21 Anterior end not transformed into a tentacular crown (NOTE: Antennae and tentacular cirri may be crowded near the anterior end) 25 Tube calcareous; thoracic membrane present 22 Tube raucoid or homy, often covered with sand-grains; thoracal membrane absent 23 Tube irregularly twisted or straight, sometimes coiled near base; body symmetrical; more than four thoracic setigers SERPULIDAE
g76 22b (21a). 23a (21b). 23b (2lb). 24a (23a). 24b (23a). 25a (20b). 25b (20b). 26a (25b). 26b (25b). 27a (26a).
27b (26a).
28a (27b). 28b (27b). 29a (26b). 29b (26b). 30a 30b 31a 3 lb 32a 32b (29a). (29a). (30a). (30a). (30b). (30b).
33 a (32a). 33b (32a). 34a (33b). 34b (33b). 35a (34a). 35b (34a). 36a 36b 37a 37b 38a 38b 39a 39b (34b). (34b). (36a). (36a). (37b), (37b). (38b). (38b).
Tube completely coiled; body asymmetrical; four thoracic setigers present SPIRORBIDAE (Usually considered a sub-family of the SERPULIDAE) Parapodia with uncini in one or a few distinct rows; tentacular crown with smooth or pennate radiles 24 Small uncini massed in dense fields in the neuropodia only; short tentacular crown with branching tentacles OWENIIDAE (part) Digestive tract recurved with anus far anteriorly; thorax without hooks except in the first setiger CAOBANGIIDAE Digestive tract straight with far posterior or terminal anus; thorax with hooks in most setigers SABELLIDAE Setiger 4 with one or a few thick spines; some median parapodia strongly modified, usually fanshaped; tubes parchmentlike, or, if homy, distinctly annulated CHAETOPTERIDAE Setiger 4 without thick spines (NOTE: Other setigers may have modified setae); no parapodia fanshaped; tubes never parchmentlike, if homy, then without annulations 26 Numerous tentacles on the lower side of the prostomium or on the peristomium; branchiae, if present, limited to a few pairs of anterior setigers 27 Anterior end with a limited (10 or fewer pairs, usually) number of antennae and tentacular cirri, or without appendages 29 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) 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 Thoracic uncini long-handled, abdominal ones short-handled TRICHOBRANCHIDAE Both thoracic and abdominal uncini short-handled; sometimes with a posterior prolongation in thoracic uncini TEREBELLIDAE Prostomium with at least one pair of antennae; peristomium usually with paired palps or tentacular cirri 30 Prostomium without appendages or with a single antenna; peristomium with paired dorsal palps, maximally two pairs of tentacular cirri or without appendages 58 Prostomium continued posteriorly in a caruncle; large notosetae furcate; others smooth or serrated . 31 Caruncle absent; furcate notosetae, if present, small, or furcate setae only kind of setae present . 32 Notosetae arranged in transverse rows on dorsum; branchiae shorter than setae . EUPHROSINIDAE Notosetae in tufts on the notopodial lobes; branchiae conspicuous branching tufts AMPHINOMIDAE Palps absent 33 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 Setae absent; acicula present only in the prolonged acicular lobes of the second segment (first segment in juveniles); otherwise absent TOMOPTERIDAE Setae or acicula or both present in most segments 34 Prostomium long and conical; usually annulated, with two pairs of antennae at the tip 35 Prostomium no more than twice as long as wide, never annulated; antennae long or short 36 Eversible pharynx with four jaws in a cross; parapodia either all uniramous or all biramous GLYCERIDAE Eversible pharynx with more than four jaws; parapodia uniramous anteriorly and biramous posterioriy GONIADIDAE Jaws present 37 Jaws absent 40 Each Jaws consisting of a series of denticles in a row DORVILLEIDAE (part) Each Jaw consisting of a single piece 38 Four or five paii of jaws LYSARETIDAE A single pair of jaws present 39 Both composite and simple setae present, parasitic in decapod crustaceans IPHITIMIDAE All setae composite, parasitic in fish ICHTHYOTOMIDAE
^ 40a 40b 41 a 41b (38b). (36b). (40b). (40b).
Science Series 28
42a (41b). 42b (41b). 43a (42b). 43b (42b). 44a (43b). 44b 45a 45b 46a (43b). (44b). (44b). (45b).
46b (45b). 47a (46b). 47b (46b). 48a (47b). 48b (47b). 49a (32b). 49b (32b). 50a (49a). 50b (49a). 51a (50b). 51b (50b). 52a (49b). 52b (49b). 53a (52b). 53b (52b). 54a (53a). 54b (53a). 55a (54b). 55b (54b). 56a (55b). 56b (55b). 57a (53b). 57b (53b). 58a (29b). 58b (29b). 59a (58a). 59b (58a).
One pair of antennae; interstitial forms PROTODREJDAE More than one pair of antennae 4j Eyes larger than the rest of the prostomium, with well defined lenses and pigment layers ALCIOPIDAE Eyes, if present, smaller than the prostomium proper, usually small pigment cups, but lensed eyes occur 42 Series of large epithelial capsules on the dorsum; two or three pairs of lateral antennae and one median, unpaired antenna SPHAERODORIDAE Epithelial capsules absent (NOTE: Etorsal cirri of phyllodocids may be inflated, but are associated with the parapodia rather than situated on the dorsum) 43 Dorsal cirri large and folise PHYLLODOCIDAE Dorsal cirri, if present, ciniform 44 Setae absent; with a few pairs of appendages anteriorly and posteriorly, parasitic in lobsters HISTRIOBDELLIDAE Setae present; several pairs of appendages along the body 45 All setae composite, pelagic LOPADORHYNCHIDAE At least some setae simple, benthic or parasitic forms 46 With ciliary bands on the dorsum of each segment in addition to a large ciliary organ on the prostomium; minute interstitial forms NERILLIDAE External ciliation limited mostly to small patches in sensory and respiratory organs, small or large forms 47 Five prostomial antennae, inquilines in bivalved moUusks ANTONBRUUNIDAE Four antennae, free-living benthic forms 48 Interramal ciiri between the noto- and neuropodia in most forms, all setae simple .. NEPHTYIDAE Interramal cirri absent; notosetae simple, neurosetae composite LACYDONIIDAE Palps bi- or multiarticulated 50 Palps simple, sometimes fused to the prostomium so the latter appears cleft or forming ventrolateral pads on the peristomium 52 Palps multiarticulated; tentacular cirri absent DORVILLEIDAE (part) Palps biarticulated; at least one pair of tentacular cirri 51 Pharynx with paired jaws; paragnaths or soft papillae or both on the surface of the everted pharynx or pharynx smooth, parapodia usually biramous NEREIDAE Pharynx usually without jaws; paragnaths or pharyngeal papillae always absent; everted pharynx often with a circlet of distal papillae or lappets, parapodia often sub-biramous or uniramous HESIONIDAE Palps ventrolateral pads on the peristomium; five occipital and two frontal antennae ONUPHIDAE (part) Palps either fused anteriorly to the prostomium or as free ventrolateral projections; maximally five antennae 53 Palps free ventrolateral projections, sometimes fused to each other 54 Palps fused to the prostomium so that the latter appears anteriorly cleft 57 Prostomium longer than wide, with a pair of antennae at the tip PISIONIDAE (part) Prostomium no longer than wide (NOTE; Fused palps may make it appear longer than it is) 55 Jaws present DORVILLEIDAE (part) Jaws absent 56 Parapodia strongly prolonged, with long supportive acicula; anterior part of the digestive tract not visibly separated into distinct parts, pelagic PONTODORIDAE Parapodia not strongly prolonged; acicula short; anterior part of the digestive tract with a visible proventriculus in most species SYLLIDAE Eversible pharynx, if present, unarmed PILARGIIDAE Eversible pharynx with four pairs of upper and one pair of lower jaws EUNICIDAE Anterior end, including both pro- and peristomium without appendages (NOTE: Appendages may be present on some anterior setigers) 59 Prostomium with a single median antenna, or peristomium with paired palps or tentacular cirri or both pro- and peristomium equipped as indicated 85 Paired palps on the first or one of the first postperistoraial segments 60 Paired palps absent 61
1976 60a (59a). 60b (59a). 61a (59b). 61b (59b). 62a (61b). 62b (61b). 63a (62b). 63b (62b). 64a (63a). 64b (63a). 65a (64b). 65b (64b). 66a (65b). 66b 67a 67b 68a 68b 69a (54b). (66b). (66b). (63b). (63b). (68a).
13
69b (68a). 70a (68b). 70b (68b). 71a (70a). 71b (70a). 72a (71b). 72b (7lb). 73a 73b 74a 74b 75a 75b 76a (70b). (70b). (73a). (73a). (74b). (74b). (73b).
76b (73b). 77a (76b). 77b (76b). 78a (77b). 78b (77b). 79a (78b).
Both composite faicigers and simple setae present ACROCIRRIDAE (part) All setae simple, distally curved or straight, sometimes acicular CIRRATULIDAE (part) With a single mid-dorsal palp on one of the first setigers (setigers 3-6 usually) .... COSSURIDAE Mid-dorsal palp absent 62 With series of long, slender branchial filaments and tentacular and dorsal cirri along the body (often lost, scars remain) CIRRATULIDAE (part) Branchial filaments and tentacular cirri absent or limited to a few segments 63 Parapodia strongly reduced so that the setae appear sessile on the body-wall; all setae simple, true capillary setae absent 64 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 Thorax with series of long dorsal cirri supported by acicula PSAMMODRILIDAE Body usually not clearly regionated; apart from papillae and reduced parapodia, other appendages absent 65 At least some setae with an internal structure of small chambers FLABELLIGERIDAE (part) No setae chambered 66 With a papilla between the rami of each parapodium; setae smooth and slightly recurved FAUVELIOPSIDAE Without papillae between the rami; setae otherwise 67 Setae in four fascicles on each segment (biramous condition) CTENODRILIDAE Setae in two fascicles on each segment (uniramous condition) PARERGODRILIDAE Prostomium an oblique plaque, usually bordered by a flange 69 Prostomium pointed, rounded or blunt 70 Setae include anterior spines, rostrate long-shafted uncini and spinose or smooth capillaries; segments usually elongated (bamboo-worms) MALDANIDAE Setae include bilimbate and spatulate kinds, long-handled uncini and companion-setae; segments not prolonged SABELLONGIDAE 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 Body not separated into regions; setal distribution and parapodial shapes grade along the body .. 73 Thorax with lateral parapodia, abdomen with both noto- and neuropodia in dorsal positions ORBINIIDAE Parapodia lateral in all parts of the body; notopodia often reduced in posterior segments and neuropodia may form nearly complete cinctures 72 Slender capillary setae in thorax and sometimes in the first few abdominal segments only; branchiae, if present, retractable filaments on the abdomen CAPITELLIDAE Slender capillary setae present on anterior, median and sometimes posterior parts of the body, including the branchial region; branchiae non-retractable, bushy or simple filaments ARENICOLIDAE Anterior end with complex jaw-apparatus 74 Jaw-apparatus absent 76 Each jaw consisting of a series of small denticles DORVILLEIDAE Each jaw consisting of a single piece 75 Hooded hooks present in at least some setigers; one pair of maxillary carriers .. LUMBRINERIDAE Hooded hooks absent; three maxillary carriers ARABELLIDAE Branchiae present on maximally 15-20 segments, starting from one of setigers 4-10 PARAONIDAE (part) Branchiae, if present, either limited to the extreme anterior end, or found scattered over a large part of the body 77 Setae include anterior spines, rostrate long-shafted uncini and spinose or smooth capillaries; segments usually elongated (bamboo-worms) MALDANIDAE Setal distribution otherwise; segments rarely elongated 78 Setae include simple capillaries and simple bifid faicigers in both noto- and neuropodia in a long region of the body QUESTIDAE Simple bifid faicigers absent 79 Prostomium an elongated cone, usually more than twice as long as wide, nearly always articulated 80
14 79b (78b). 80a (79a). 80b (79a). 81a 81b 82a 82b 83a 83b 84a (79b). (79b). (81b). (81b). (82b). (82b). (83b).
Science Series 28
84b (83b). 85a (58b). 85b (58b). 86a (85a). 86b 87a 87b 88a 88b 89a 89b 90a 90b 91a 91b 92a 92b 93a 93b 94a 94b (85a). (86b). (86b). (85b). (85b). (88a). (88a). (88b). (88b). (90b). (90b). (91b). (91b). (92b). (92b). (93b). (93b),
Prostomium less than twice as long as wide, may be bluntly conical, rounded or truncate, never articulated 81 Eversible pharynx with four jaws; parapodia either all uniramous or all biramous GLYCERIDAE (part) Eversible pharynx with more than four jaws; anterior parapodia uniramous, posterior ones biramous GONIADIDAE Parapodia rounded lobes with large, easily dehiscent cirri; pelagic TYPHLOSCOLECIDAE Parapodia low folds or blunt, button-shaped projections; cirri, if present, cirriform 82 Composite falcigers present in anterior setigers ACROCIRRIDAE (part) All setae simple 83 Neurosetae uncinal in structure BOGUEIDAE Uncini absent 84 AH setae simple capillaries; branchiae cirriform, pectinate or absent; prostomium entire, pointed or rounded OPHELIIDAE Furcate and acicular setae usually present; branchiae, if present, limited to the anterior end and strongly arborescent; prostomium T-shaped or bifid SCALIBREGMIDAE Prostomium with a median antenna 86 Prostomium without appendages 88 Branchiae present on maximally 15-20 segments first starting from one of setigers 4-10 PARAONIDAE (part) Branchial distribution otherwise or branchiae absent 87 Notopodial cirri flask-shaped in some setigers, plumose setae present POECILOCHAETIDAE Notopodial cirri cirriform or folise, plumose setae absent SPIONIDAE (part) With two pairs of tentacular cirri; setae composite 89 Without tentacular cirri; setae simple 90 Setal appendages long and slender; pelagic lOSPILIDAE Setal appendages short; parasitic CALAMYZIDAE With multiple series of small, long-shafted uncini in the neutopodia OWENIIDAE (part) Uncini in single rows or absent 91 Prostomium flattened and spatulate; as wide as the widest part of the body MAGELX)NIDAE I*rostomium not flattened; distinctly narrower than the widest part of the body 92 Parapodia inconspicuous; abdominal segments prolonged with setae forming complete cinctures around the body HETEROSPIONIDAE Parapodia well developed; abdominal segments usually not prolonged; setae limited to lateral tufts 93 All parapodia biramous, except possibly the first one 94 Median parapodia uniramous TROCHOCHAETIDAE Uncini absent; notopodia with acicula, but without setae; true branchiae absent APISTOBRANCHIDAE 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 asetigerous anterior segments present; no additional cephalized segments present. Palps absent; eversible pharynx either an axial sac or a ventrolateral pad. Parapodia biramous; all setae simple, including capillary setae and usually acicular spines and serrated or spinose setae.
FAMILY ORBINIIDAE HARTMAN
1942
Orbiniida with lateral parapodia in a thoracic region and, usually, dorsal parapodia in an abdominal region.
Prostomium without appendages, one or two asetigerous segments present anteriorly; no peristomial appendages. A saclike proboscis present. All setae simple, including capillaries, simple hooks and sometimes brush-topped, bifid or furcate setae. The orbiniids have been the subject of several comprehensive studies, including Hartman (1957), Pettibone (1957a) and Day (1973). The major generic groupings appear clear and were reviewed by Day (1973) for the subfamily Orbiniinae. The subfamily Protoariciinae has yet to be reviewed in detail.
1976
15
FIGURE
2. (A), Family ORBINHDAE, Orhinia johnsoni, anterior end, after Hartman 1969, 5x; (B), transverse section of the abdomen of the above, 18x; (c). Family PARAONIDAE, Cirrophorus sp., anterior end, diagrammatic, about 22x; (D), Family QUESTIDAE, Quesia caudicirra, posterior seta, after Hartman 1966, 54x; (E) anterior end of the above, 24x.
KEY TO GENERA
la. lb. 2a (la). 2b (la). 3a (2b). 3b (2b). 4a (3b). 4b (3b). 5a (4b). 5b (4b). 6a (5b). 6b (5b). 7a (lb). 7b (lb). 8a (7b). 8b (7b). 9a (8b). 9b (8b). 10a (9a).
Two asetigerous anterior segments A single asetigerous anterior segment Branchiae absent Branchiae present Branchiae present on all but a few anterior segments Branchiae limited to abdominal segments Only crenulated capillaries present Crenulated capillaries and other kinds of setae present Abdominal neurosetae include swan-shaped hooks Swan-shaped hooks absent Abdominal neurosetae all crenulated capillaries; prostomium rounded Abdominal neurosetae include acicular hooks; prostomium pointed Branchiae absent Branchiae present on at least some segments Prostomium rounded or truncate Prostomium more or less pointed All thoracic parapodia with only slender, pointed setae Some thoracic neuropodia with setae of another kind Abdominal neuropodia with acicular spines
2 7 Orbiniella 3 Protoariciella 4 Scoloplella 5 Proscoloplos 6 Protoaricia Schroederella Microrbinia 8 Naineris 9 10 11 Berkeleya
16 10b (9a). 1 la (10b). 1 lb (10b). 12a (1 la). I2b (11a). 13a (lib). 13b (I lb). 14a (13b). 14b (13b).
Science Series 28
Acicular spines absent in abdominal neuropodia Haploscoloplos Thoracic neuropodia of two abruptly different kinds 12 Thoracic neuropodia not abruptly different 13 Anterior three thoracic neuropodia with bristle-tipped setae Califia Posterior thoracic neuropodia with thick, modified spines associated with a glandular pouch , . Phylo Some thoracic segments with rows of papillae along the ventrum, papillae sometimes also on parapodial postsetal ridges Orbinia Without rows of papillae on the ventrum 14 Median and posterior abdominal neuropodia with thick acicula, projecting from the parapodial lobes; branchiae usually present from fifth or sixth segment Scoloplos (Leodamas) Thick projecting acicula absent; branchiae usually not present before tenth segment (Scoloplos)
Generic Definitions Berkeleya Hartman 1971, B. profunda Hartman 1971; only species. ORBINIINAE with pointed prostomium, abdominal branchiae and all thoracic setae crenulated capillaries. Twelve thoracic segments; transition from thorax to abdomen abrupt. Abdominal notopodia with crenulated capillaries and furcate setae; abdominal neuropodia with crenulated capillaries and acicular spines. Califia Hartman 1957, C, candida Hartman 1957; 4 species. ORBINIINAE with pointed prostomium, branchiae not present before segment 8; posterior thoracic neuropodia with two or fewer papillae; ventral papillae absent. Thoracic neurosetae include anterior brush-topped setae, crenulated capillaries and in some cases blunt hooks. Haploscoloplos Monro 1933, Scoloplos cylindrifer Ehlers 1905; 13 species. ORBINIINAE with pointed prostomium, branchiae not present before setiger 9; posterior thoracic neuropodia with two papillae and maximally two ventral papillae present; never more than four papillae combined. All neurosetae crenulated capillaries; notosetae crenulated capillaries and in some cases furcate setae. Microrbinia Hartman 1965, Ai. linea Hartman 1965; only species. ORBINIINAE with conical prostomium; branchiae and accessory papillae absent. Separation between thorax and abdomen indistinct. First notosetae smooth capillaries; other notosetae camerated capillaries and posterior notosetae curved and serrated acicular spines. Neurosetae similar, Naineris Blainville 1828, Nais quadricuspida Fabricius 1780; 18 species. ORBINIINAE with rounded or truncate prostomium, branchiae first present from any segment from 2-23. Maximally two accessory papillae on posterior parapodia, ventral papillae absent. Thoracic neurosetae
include crenulated capillaries, hooks and intermediate forms. Abdominal notosetae crenulated capillaries and sometimes furcate setae. Orbinia Quatrefages 1865, Aricia cuvierii Audouin and Milne Edwards 1833c; 16 species. ORBINIINAE with pointed prostomium and first pair of branchiae on setiger 5-9. Posterior thoracic parapodia with several accessory papillae and numerous ventral papillae; with a combined total of at least five papillae on each segment. Thoracic neurosetae include hooks (or subuluncini) and crenulated capillaries. Furcate and capillary setae present in abdominal notopodia. Orbiniella Day 1954, O. minuta Day 1954; 4 species. PROTOARICIINAE with rounded or pointed prostomium and without branchiae. Thoracic setae all camerated or crenulated capillaries; abdominal setae include crenulated capillaries, acicular setae and sometimes furcate setae. Phylo Kinberg 1866b, P. felix Kinberg 1866b; 19 species. ORBINIINAE with pointed prostomium and branchiae first present from setiger 5-7. Posterior thoracic parapodia with several accessory papillae; numerous ventral papillae present. At least five papillae in combined total present on a segment. Thoracic neurosetae include crenulated capillaries, heavy hooks and heavy spearshaped setae. Proscoloplos Day 1954, P. cygnochaetus Day 1954; 2 species. PROTOARICIINAE with rounded prostomium and stout branchiae present from setiger 8. Transition from thorax to abdomen indistinct. Setae include crenulated capillaries and one or two swan-shaped hooks in posterior neuropodia. Protoaricia Czerniavsky 1881a, Aricia oerstedii Claparde 1864; 2 species. PROTOARICIINAE with rounded prostomium and branchiae limited to abdominal segments. Thoracic
1976
77
Invalid Genera
neurosetae include crenulated capillaries, hooks and subuluncini; abdominal neurosetae all crenulated capillaries. Protoariciella Hartmann-Schrder 1962, P. uncinata Hartraann-Schider 1962; 3 species. PROTOARICIINAE with pointed prostomium and branchiae present on all but a few anterior segments (from setiger 3 in type species). Thoracic notosctae all are crenulated capillaries; acicular setae present in posterior notopodia. Neurosetae include thick hooks with three teeth; slender acicular setae with flat teeth, crenulated capillaries and thick smooth spines in posterior segments. Schroederella Laubier 1962, S. pauliana Laubier 1962; 2 species. PROTOARCIINAE with strongly pointed prostomium and branchiae present on the abdomen, Notosetae include crenulated capillaries and acicular spines in far posterior segments. Neurosetae include crenulated capillaries and acicular hooks. Scoloplella Day 1963, S. capensis Day 1963; only species. PROTOARICIINAE with pointed prostomium and branchiae present on the abdomen. All setae crenulated capillaries. Scoloplos Blainville 1828, Lumbricus armiger 0,F. Mller 1776; 38 species. ORBINIINAE with pointed prostomium and branchiae first present from setiger 5 or later. Accessory papillae never exceeding four, including both ventral and parapodial kinds. Thoracic neurosetae include blunt hooks and crenulated capillaries (may be absent). Abdominal notosetae include crenulated capillaries, furcate setae and spines. Subgenus Scoloplos: Species with first branchiae on setigers 8-10 or later; without emerging notacicula in posterior setigers. Subgenus Leodamas: Species with first branchiae on setiger 5-6; with heavy emerging notacicula in posterior setigers.
Alcandra Kinberg 1866b, sec Scoloplos (Leodamas) Anthostoma Schmarda 1861, seeNaineris Aricia Savigny 1820, seeOrbiitia Branchetus Chamberlin 1919c, see Scoloplos (Leodamas) Clytie Grube 1855, indeterminable Gisela Mller 1858, indeterminable Labotas Kinberg 1866b, see Scoloplos Lacydes Kinberg 1866b, see Naineris Naidonereis Malmgren 1867, error for Naineris Theodisca Mller 1858, see Naineris Theostoma Eisig 1914, s&e Protoaricia Venadis Castelnau 1842, see Orbinia Incertae Sedis Falklandiella Hartman 1967, F. annulata Hartman 1967; only species. Resembles the orbiniids in setal structures, but appears to differ in other features, Prostomium a simple triangular lobe; two asetigerous segments present. Parapodia biramous with very strongly reduced parôdial lobes. Notosetae all slender and camerated; most neurosetae similar, but shorter; in addition heavy acicular spines present in posterior neuropodia.
FAMILY PARAONIDAE CERRUTI
1909
Body long and slender with lateral parapodia. Prostomium with a single antenna or antennae absent. Branchiae present on a limited number of median setigers in most species. All setae simple, including capillaries and various, usually postbranchial, hooks or otherwise modified setae. The key given below is based on the important revision by Strelzov (1973); some features are taken from Fauchald (1972). Strelzov recognized only two genera with prostomial antennae, Cirrophorus and Aricidea; the latter with a series of subgenera; these subgenera are here considered distinct genera.
Key to Genera la. lb. 2a (lb). 2b (lb). 3a (2b). 3b (2b). 4a (3a). 4b (3a). 5a (4b). Setae present 6om the first segment Aparaonis Setae present from the second segment 2 Modified setae notopodial Cirrophorus Modified setae, if present, neuropodial 3 Prostomium with a median antenna 4 Prostomium without a median antenna 7 Modified setae absent Aedicira Modified setae present 5 Modified setae either pseudocomposite or curved with a subterminal arista on concave side of the shaft Aricidea
8 5b (4b). 6a (5b). 6b (5b). 7a (2b). 7b (2b). 8a (7b). 8b (7b). 9a (8b). 9b (7b).
Science Series 28
Modified setae otherwise 6 Modified setae not greatly different from the capillary setae, but somewhat thicker and more abruptly tapering and with a long arista Allia Modified setae distally strongly curved, or bifid, either with numbers of long aristae or bearded .... Acesta Nuchal organ on posterior part of prostomium, prostomium with several transverse bands of cilia Paraonis Nuchal organ on the reduced peristomium; with a single preoral ciliary band or prostomial ciliary bands absent 8 Modified setae absent Paraonella Modified setae present 9 Three prebranchial setigers; terminal prostomial sense organs absent Sabidius More than three prebranchial setigers; terminal prostomial sense organs present Tauberia
Generic Definitions Acesta Strelzov 1973, Aricidea catherinae Laubier 1967a; 10 species. Median antenna present. Modified setae neuropodial; each strongly curved distally, or bifid with several aristae or with a short stiff beard. Aedicira Hartman 1957, Aricidea pacifica Hartman 1944c; only species. Median antenna present. All setae slender capillaries; those in the neuropodial fascicles often shorter than the notopodial ones. Allia Strelzov 1973, Aricidea albairossae Pettibone 1957b; 13 species. Median antenna present. Modified setae slightly thicker than the non-modified capillary setae; somewhat more abruptly tapering; or tapering abruptly near the middle, giving the outer part of the seta the appearance of a long, smooth arista. Aparaonis Hartman 1965, A. abyssalis Hartman 1965; only species. Setae present in the first segment; three pairs of branchiae on segments 2-4. All setae simple capillaries. Aricidea Webster 1879b, A. fragilis Webster 1879b; 7 species. Median antenna present; modified setae either pseudocomposite or recurved with a subterminal arista arising from the concave side of the shaft. Cirrophorus Ehlers 1908, C. branchiatus Ehlers 1908; 10 species. Median antenna present or absent; modified setae present in the postbranchial notopodial fascicles. Paraonella Strelzov 1973, Paraonides nrdica Strelzov 1968a; 4 species. Nuchal organs on the peristomium; median antenna absent. Modified setae absent. Paraonis Cerruti 1909, Aonides fulgens Levinsen 1884; 2 species.
Nuchal organs prostomial; median antenna absent. Modified setae neuropodial. Sabidius Strelzov 1973, Paraonis cornatus Hartman 1965; only species. Nuchal organs peristomial; median antenna absent. Modified setae neuropodial. Prostomial terminal sense organs absent. Three prebranchial setigers. Tauberia Strelzov 1973, Aonides gracilis Tauber 1879; 10 species. Nuchal organs peristomial; median antenna absent. Modified setae neuropodial. Prostomial terminal sense organs present. More than three prebranchial setigers present. Taxonomic Notes The generic subdivision given above, differs markedly from the more traditional pattern summarized by Fauchald (1972). Strelzov (1973) introduced several new taxonomic characters to the ones previously in common usage. The presence of prostomial ciliary bands, of prostomial terminal sense organs and the localization of the nuchal organs must now be investigated. Similar characters have been used in other famihes, and it is anticipated that they will gain acceptance also in the treatment of the paraonids. Invalid Genus Levinsenia Mesnil 1897, see Paraonis. Several other genera, such as Paraonides. Paradoneis and others used in the traditional sub-division of the family, may be invalid. Until the value of Strelzov's system has been tested, I hesitate to treat these genera as invalid.
FAMILY QCIESTIDAE HARMAN
1966a
Body long and slender. Prostomium without appendages. One asetigerous anterior segment present. Pharynx with a muscular pad. Parapodia biramous with reduced lobes. Branchiae and anal cirri present or
1976
\9
absent. Setae include serrated capillaries, bidentate hooks and trifrcate spines. As remarked by Hobson ( 1970) the questids have the gonads Umited to a few segments; this generally is considered the key feature separating the oligochaetes
from the polychaetes (Fauchald 1974a). However, the questids have several different kinds of setae; a feature rarely found among the oligochaetes. It is quite possible that the family should be considered among the oligochaetes.
Key to Genera la. lb. Setae include capillaries and bidentate hooks; posterior cirriform branchiae present Setae include capillaries, bidentate hooks and trifrcate spines; branchiae absent Generic Definitions Novaquesta Hobson 1970, N. trifitrcata Hobson 1970; only species. Questids with serrated capillary setae, bidentate hooks and trifrcate spines with the middle tine shorter and slenderer than the other; branchiae and anal cirri absent. Questa Harman 1966a, Q. caudicirra Hartman 1966a; only species. Questids with serrated capillary setae and bidentate hooks. Branchiae present on posterior segments, anal cirri present. ORDER CTENODRILIDA Prostomium without appendages; palps absent. At least one asetigerous anterior segment present. Proboscis a ventral muscular pad. Parapodia uni- or biramous; all setae simple. Parapodial lobes absent. Members of this order are very small, generally grub-shaped polychaetes that tend to turn up in massculture in aquaria more frequently than in the field (especially true for the ctenodrilids). Specimens of Ctenodrilus also have turned up associated with FlabelUderma commensalis at Santa Catalina Island and have the same dark purple pigmentation on sea urchins as these commensals do.
FAMILY CTENODRILIDAE KENNEL
Quesxa Novaquesta
(1971); it now is divided into two sub-families with three genera in all; each with only a few species.
1882
Body short or long, prostomium without appendages, palps absent. Pharynx a ventral muscular pad. Some anterior segments may be fused and asetigerous; some segments with median dorsal cirri; paired lateral branchiae present in some juveniles. Setae in double fascicles; parapodial lobes absent. This small family of tiny polychaetes recently has been reorganized and redefined by Hartmann-Schrder
FIGURE 3. (A), Family CTENODRILIDAE, Ctenodrilus serratus, modified after Hartman 1944, 284x; (B), Family PARERGODRILIDAE, Stygocapitella subterrnea, after HanmannSchrder 1971, about 30Ox.
Key to Genera la. lb. Body short and gniblike, with few segments, unpaired dorsal cirri and branchiae absent CTENODRILINAE Ctenodrilus Body long and slender, with at least 15 segments; unpaired dorsal cirri on one or two anterior segments or larval branchiae present RHAPHIDRILINAE 2
20 2a (lb). 2b (lb).
NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY Unpaired dorsal cirri present; branchiae absent Unpaired dorsal cirri absent; branchiae present Generic Definitions
Science Series 28 Zeppelinia Raphidrilus
Ctenodrilus Claparde 1863, Partkenope cirrata Schmidt 1857; 2 species, CTENODRILINAE. Body with maximally 15 segments; one achaetous segment present. Setae thick, marginally coarsely dentate or smooth. Rkapkidrilus Monticelli 1910, , nemasoma Monticelli 1910; only species, RHAPHIDRILINAE. Body long and slender, with at least 18 segments; one asetigerous segment present. Setae include capillaries and genital spines in the adult males. Paired branchiae present in the juvenile stages, Zeppelinia Vaillant 1890, Ctenodrilus monostylos Zeppelin 1883; 5 species. RHAPHIDRILINAE. Body long and slender, with at least IS segments; asetigerous segments absent.
Setae include capillaries and short, thicker dentate or smooth spines. One or two unpaired dorsal cirri present anteriorly. Invalid Genera Monostylos Vejdovsky 1884, see Zeppelinia Partkenope Schmidt 1857, set Ctenodrilus
FAMILY PARERGODRILIDAE REISINGER
1960
Small, grub-shaped worms with at least one anterior asetigerous segment. Prostomium without appendages; parapodial lobes absent; setae present in single fascicles, each with one or a few setae only. Setae all simple. The parergodrilids are very small worms, Uving either interstitially in shallow marine sands (Stygocapitella) or in rotting terrestrial plant material (Parergodrilus). The family is poorly known and understood; the revision by Hartmann-Schrder (1971) is followed here.
Key to Genera la. lb. Body not translucent, with 10-11 setigers; setae limbate and furcate Body translucent, 8-9 setigers present; all setae smooth rods Generic Definitions Parergodrilus Reisinger 1925, P. keideri Reisinger 1925; only species. One anterior and no posterior asetigerous segments; eight or nine setigers present. Setae all smooth rods, except copulatory hooks in males. Stygocapitella Knllner 1934, S. subterrnea KnUner 1934; only species. Two anterior and two posterior asetigerous segments, ten or 11 setigers present. Setae include smooth, short bilimbate setae, smooth bilimbate setae with long, whiplike tips and short furcate setae with two accessory teeth in the crotch. ORDER PSAMMODRILIDA Prostomium without appendages, palps absent. At least one asetigerous anterior segment present. Notopodia in a median region strongly prolonged and supported by acicula, Parapodia biramous; neuro|X)dial tori present in one form, otherwise parapodial lobes absent (except for the cirri mentioned above); all setae simple.
FAMILY PSAMMODRILIDAE SWEDMARK
Stygocapitella Parergodrilus
1952
FiGimE 4. (A), Family PSAMMODRILIDAE, Psammodriloides fauveli after Swedmark 1958, about 50x; (B), Family CO%S\JRJDk'E,Cossurabrmnea, afterFauchald 1972, 12.5x,
Small, grub-shaped worms. Pro- and peristomium without appendages; palps absent. Parapodia, except
]97()
2/
the thoracal dorsal cirri reduced; abdominal neuropodial tori present in one form. As remarked by Swedmark (1958), these small interstitial worms are very isolated among the polychaetes.
The first form described, Psantmodritus balanoglossoides Swedmark, is unique in that the eversible pharynx is formed from the longitudinal body muscles.
Key to Genera la. lb. Dorsal cirri decreasing evenly in length from the first to the sixth thoracic segment; abdominal uncinigers with several uncini Psammodrilus 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 Psammodriloides Swedmark 1958, P. fauveli Swedmark 1958; only species. Body with two distinct regions: head and trunk; six thoracic setigers, each with a dorsal cirrus supported by an aciculum. Ten abdominal setigers; each with a single uncinus in each neuropodium. Eversible pharynx absent. Psammodrilus Swedmark 1952, P. balanogtossoides Swedmark 1952; only species. Body in three regions; head, pharyngeal region and trunk. Six thoracic setigers with dorsal cirri supported by acicula; up to about thirty abdominal setigers; each with several uncini in each neuropodium. Pharyngeal apparatus present. ORDER COSSURIDA Prostomium without appendages; a single peristomial asetigerous segment present; a single median palp present on the dorsum of an anterior setiger (usually between setigers 3 and 6). Proboscis a ventral pad. Parapodia biramous, with very low parapodial lobes; all setae simple.
FAMILY COSSURIDAE DAY
rata Webster and Benedict (1887) with a total of about fifteen species. Two of these differ rather sharply from the others and may deserve separate generic standing. ORDER SPIONIDA Prostomium distinct, without appendages or with an occipital antenna, A pair or two groups of grooved feeding palps usually present anteriorly, either on the prostomium or on an anterior segment. Paired tentacular cirri sometimes present at the base of the palps. Pharynx either axial or as a ventral pad, Parapodia well developed or reduced. All setae simple.
1963
Prostomium without appendages, a single median palp present dorsally on one anterior setiger; proboscis a ventral pad. Parapodia biramous, with reduced parapodial lobes. All setae simple, including bilimbate or hirsute setae in two or more fascicles; thick spines and capillary setae present in the abdomen of some forms. Cossurids are common in sand and especially in deep slope and abyssal muds. They are burrowers, and apparently feed on detritus with the help of the pharynx. The palp in this case appears to be sensory, and perhaps may additionally be respiratory in function since it is well-equipped with blood-vessels (Fauchald in preparation). The only currently described genus is Cossura Webster and Benedict, (1887) with genotype C. longocir-
FIGURE 5. (A), Family APISTOBRANCHIDAE, Apistobranchus tullbergi, ventral view, after Pettibone 1963, lOx; (B), dorsal view of the above, lOx; (C), Family TROCHOCHAETIDAE, Trochochaetus multisetosum, lateral view, after Pettibone 1963, lOx; (D), dorsal view of the above, lOx; (E), Family SPIONIDAE, Boccardia proboscidea, modified after Hartman 1969, 25x.
22
NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY Key to Suborders
Science Series 28
la. lb. 2a (lb). 2b (lb).
Body separated into at least two distinct regions; uncini present Chaetopterifonnia Body rarely separated into distinct regions, uncini always absent 2 Palps at the junction of the pro- and peristomium; parapodial lobes well developed at least in some setigers Spioniformia Palps post-peristomial in origin; parapodial lobes poorly developed Cirratuliformia Suborder Spioniformia
FAMILY SPIONIDAE GRUBE
1850
Palps on the peristomium or at the junction between the pro- and peristomium; eversible pharynx either a ventral pad or an axial sac.
FAMILY APISTOBRANCHIDAE MESNIL AND CAULLERY
1898
Spioniforms with the anterior end inflated and the rest of the body cylindrical. Prostoraium without appendages, but with palps attached at the postectal margins. Parapodia biramous, parapodial lobes mostly cirriform, but serrated postsetal lobes present in some setigers. All setae simple, mainly capillaries. The apistobranchids are known for one genus, Apistobranchus Levinsen 1883, with type species Aricia tullbergi Theel 1879, A total of three species currently are recognized. Apistobranchids are in fact not as rare as the low number of species should indicate; they are, as some of their spionid relatives, not limited to a tubicolous existence, but will feed from loosely constructed burrows. Most commonly found in shelly sands, sands and muds. Invalid Genera Ethocles Webster and Benedict 1887, see Apistobranchus Skardaria Wesenberg-Lund 1951, see Apistobranchus
Spioniforms with body elongated. Prostomium anteriorly blunt, with frontal horns, or pointed; an occipital papilla may be present, other appendages absent. Palps at the postectal comers of the prostomium. Parapodia biramous, parapodial lobes cirriform or folise, never serrated. All setae simple, including capillaries and bi- or multidentate, hooded or non-hooded hooks. Spionids are very common in all environments. Some forms are burrowers (Polydora, Boccardia et al.) in calcareous substrata or in rock; others build permanent tubes in soft substrata and some forms combine the two activities. Numerous forms are freeliving in sands and muds. Species of some genera, most frequently perhaps of Prionospio SENSU LAW and of the PolydoralBoccardia complex, very likely are present in any kind of benthic marine soft-bottom sample taken anywhere on the globe. Members of the genus Aonides are dominant forms in sandy beaches in temperate and warm areas of the world (Foster 1971). The spionids usually lose their palps, and frequently most of their branchiae in fixation. This is especially true in those genera where the structure of the branchiae is important for species separation. Attempts at identification of these taxa on incomplete material should be avoided; the family is speciose and serious errors are easUy introduced into the literature, if care is not taken at all identificatoiy levels.
Key to Genera
la. lb. 2a (lb). 2b (lb). 3a (2a). 3b (2a). 4a (3b). 4b (3b). 5a (4b). 5b (4b). 6a (5b). 6b (5b). 7a (6b). 7b (6b), 8a (2b).
Each branchia with several (maximally seven) processes in median parapodia Potybranchia Branchia, if present, single process 2 One setiger with strongly modified setae 3 Setae change gradually along the body, no segment with remarkably different setae (except large hooks in the first setiger) 8 Setiger 16 modified Morants Either setiger 4 or 5 modified 4 Setiger 4 modified Polydorella Setiger 5 modified 5 Neuropodial uncini distally trifid Tripolydora Neuropodial uncini distally bifid or entire 6 Branchiae first present from setiger 2 Boccardia Branchiae first present posterior to the modified segment 7 Setae in modified segment arranged in a horseshoe-shaped series Pseudopolydora Setae in the modified segment in a straight line or a small patch Polydora Branchiae absent; first neuropodia with stout hooks Spiophanes
]g7(, 8b (2b). 9a (8b). 9b (8b). 10a (9a). 10b (9a). 1 la ( 10a). lib (10a). 12a ( 1 lb). 12b (1 lb). 13a (12b). 13b (12b). 14a (10b). 14b (10b). 15a ( 14b). I5b (14b). 16a (15a). 16b (15a). 17a (15b). 17b ( 15b). 18a (17b). 18b (17b). 19a (9b). 19b (9b). 20a (19b). 20b 21a 21b 22a 22b 23a 23b 24a 24b 25a 25b 26a 26b 27a Tib (19b). (20b). (20b). (21a). (21a). (22b). (22b). (21b). (21b). (24b). (24b). (25b). (25b). (26b). (26b).
2i
At least one pair of branchiae present; first neuropodia without stout hooks 9 Branchiae present on nearly the whole body 10 Branchiae limited to less than half the length of the body 19 Branchiae present from setiger 1 11 Branchiae present from setiger 2 14 Accessory branchiae on some segments Dispio Accessory branchiae absent 12 Posterior notopodia with uncini Marenzelleria Notosetae all capillaries 13 Prostomium anteriorly rounded Spio Prostomium with laterofrontal horns Malacoceros Posterior notopodia with uncini Scolelepis All notosetae capillaries 15 Prostomium with laterofrontal horns 16 Prostomium anteriorly blunt or pointed 17 Branchiae partially fused to the notopodial postsetal lobes Rhynchospio Branchiae completely separated from the notopodial postsetal lobes Mesospio Occipital antenna absent Microspio Occipital antenna present 18 Branchiae partially fused to the notopodial postsetal lobes Pseudomalacoceros Branchiae completely separated from the notopodial postsetal lobes Laonice A single pair of branchiae present Streblospio At least two pairs of branchiae present 20 Branchiae concentrated in a medio-posterior region, except in the males, where an additional single pair is present on the second setiger Pygospio Branchiae concentrated near the anterior end only 21 Peristomium forms large lateral wings on either side of the prostomium 22 Peristomium does not form lateral wings on the sides of the prostomium 24 Three pairs of pinnate branchiae present from setiger I Paraprionospio Branchiae otherwise, first present from setiger 2 23 Two to four pairs of branchiae present Aquilaspio At least six pairs of branchiae present Minuspio Prostomium anteriorly pointed Aonides Prostomium anteriorly blunt or with frontal homs 25 Prostomium with frontal homs Scolecolepides Prostomium anteriorly rounded 26 Branchiae two pairs on setigers 34 Anaspio Four pairs of branchiae from setiger 2 27 Three first pairs of branchiae cirriform, the last pinnate Apoprionospio Branchiae cirriform or pinnate in another pattern Prionospio free from the notopodial postsetai lobes. Anterior setigers with capillary setae only; posterior parapodia with uncini in both rami. Neuropodial postsetal lobes smooth. Apoprionospio Foster 1969, A. day i Foster 1969; 5 species. Prostomium anteriorly blunt; peristomium without lateral wings. Four pairs of branchiae from setiger 2; first three cirriform, fourth pinnate. All anterior setae capillaries; posteriorly uncini in both rami. Neuropodial postsetal lobes large and folise in setiger 2. Aquilaspio Foster 1971, Prionospio sexoculata Augener 1918; 2 species.
Generic Definitions Anaspio Chamberlin 1920, A. boreus, Chamberlin 1920; only species. Prostomium anteriorly rounded. Two pairs of branchiae on setigers 3 and 4; free from the notopodial postsetal lobes. Notosetae all capillaries (in 36 setigers); neuropodia with uncini from setigers 10-11. Anterior parapodial lobes very large and folise. Aonides Claparde \%(A,Nerine oxycephala Sars 1862; 7 species. Prostomium anteriorly pointed. Branchiae from setiger 2, present on anterior end only; completely
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Prostomium anteriorly blunt; peristomium with large lateral wings; first setiger more or less reduced. Two to four pairs of branchiae from setiger 2. All anterior setae capillaries; posterior setigers with uncini in both rami. Boccardia Carazzi 1895, Polydora (Leucodore) polybranchia Haswell 1885; 15 species. Prostomium anteriorly blunt or bifid. Branchiae present from setiger 2. Fifth setiger modified with strong setae; in other setigers, all notopodia with capillaries only; neuropodia posteriorly with uncini. Dispio Hartman 1951a, D. uncinata Hartman 1951a; 5 species. Prostomium anteriorly blunt; peristomium enfolding sides of prostomium. Branchiae from setiger 1 to the end of the body; at least partially fused to the notopodial postsetal lobes; accessory branchiae present in some setigers. Notosetae all capillaries; neurosetae include capillaries and uncini in median and posterior setigers. Laonice Malmgren 1867, Nerine cirrata Sars 1850; 15 species. Prostomium anteriorly rounded; occipital antenna present. Branchiae present from setiger 2 to the middle of the body; completely free from the notopodial postsetal lobes. Notosetae all capillaries; neurosetae include capillaries and uncini. Genital pouches in at least some setigers. Malacoceros Quatrefages 1843, Spio vulgaris Johnston 1827; 10 species. Prostomium with lateral horns. Branchiae present from setiger 1, partially fused to the notopodial postsetal lobes. Notosetae all capillaries; neurosetae include capillaries and uncini in posterior setigers. Marenzelleria Mesnil 1896, M. wireni Augener 1913b; only species. Prostomium anteriorly rounded. Branchiae present from setiger 1, partially fused to the notopodial postsetal lobes. Anterior setigers with capillary setae in both rami; posteriorly uncini in both rami. Mesospio Gravier 1911a, M. moorei Gravier 1911a; only species. Prostomium with frontal horns. Branchiae present from the second setiger, completely free of the postsetal lobes. Notosetae all capillaries; neuropodia with uncini present from setiger 15. All anal cirri similar, short and tapering. Microspio Mesnil 1896, Spio mecznikowianus Claparde 1870a; 10 species. Prostomium anteriorly rounded. Branchiae present irom setiger 2; partially fused to the notopodial postsetal lobes. Notosetae all capillaries; neuropodia posteriorly also with uncini.
Minuspio Foster 1971, Prionospio cirrifera Wiin 1883; 7 species. Prostomium anteriorly blunt; peristomium with large lateral wings. Branchiae present from setiger 2 to about setiger 40, all cirriform. Anterior setae all capillaries in both rami; posterior end with uncini in both rami. Morants Chamberlin 1919a, M. duplex Chamberlin 1919a; only species. First four setigers with branchiae; fused to the notopodial postsetal lobes; setiger 16 with strongly modified setae. Curved hooks present in the first notopodia; all other notosetae capillary. Anterior neurosetae capillary, posterior ones uncini. Paraprionospio CauUery 1914b, Prionospio pinnala Ehlers 1901; 5 species. Prostomium anteriorly rounded; peristomium with large lateral wings. Second segment asetigerous. Three pairs of pinnate branchiae present from setiger 1 ; parapodia of fu^t setiger well developed. All anterior setae capillaries; posterior parapodia with imcini in both rami. Poly bronchia Potts 1928, P. foxi Potts 1928; only species. Prostomium with frontal horns. Branchia present from setiger 1; each branchia with maximally six to seven processes in median setigers. Capillary setae absent; all setae uncmi m both rami. Polydora Bosc 1802, P. cornuta Bosc 1802; 65 species. Prostomium anteriorly blunt or bifid. Branchiae not present before setiger 6. Setiger 5 with strongly modified, stout setae. Other setae include notopodial capillaries, sometimes also with simple posterior spines. Neutopodial uncini present fiom setiger 7-10 in most species. Pofydorella Augener 1914, P. prolifra Augener 1914; 2 species. Ihtjstomium anteriorly blunt. Branchiae fiwm setiger 6. Fourth setiger modified with large, stout setae. Notosetae all capillaries, except in setiger 4; neuropodia with capillaries, but with uncini present from setiger 6. Prionospio Malmgren 1867, P. steenstriipi Makngren 1867; 36 species. Prostomium anteriorly blunt; peristomium without lateral wings. Four pairs of branchiae, either cirriform or pinnate or both, first present from setiger 2. Branchial parapodia with large postsetal lobes. All anterior setae capillaries; in median and posterior parapodia uncini also present. Pseudomalacoceros Czemiavsky 1881b, Nerinides cntabra Rioja 1918; 11 species. Prostomium anteriorly blunt or pointed; occipital antenna present. Branchiae present from setiger 2; at least partially fused to the notopodial postsetal lobes. Notosetae all capillaries; neurosetae include capillaries and uncini.
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Pseudopolydora Czerniavsky 1881b, Polydora antennata Claparde 1870a; 11 species. Prostomium anteriorly blunt or bifid. Branchiae first present from setiger 7. Fifth setiger modified with large, thick setae arranged in a horseshoe-shaped pattern. Notosetae all capillaries; neurosetae include capillaries and posterior uncini. Pygospio Claparde 1863, P. ekgans Claperde 1863; 3 species. Ftestomium anteriorly rounded. Branchiae concentrated in a short posterior region, except in the males where a single pair is present on the second setiger; all are fused to the notopodial postsetal lobes. Notosetae all capillaries; neurosetae include capillaries and uncini. Rhynchospio Hartman 1936b, R. arenincola Hartman 1936b; 4 species. Prostomium with frontal horns. Branchiae present from setiger 2, partially fused to the notopodial postsetal lobes. All notosetae capillaries; neurosetae include capillaries and uncini. Scolecolepides Ehlers 1907, S. benhami Ehlers 1907; 3 species. Prostomium with frontal horns. Branchiae present fix)m setiger 1, limited to the anterior part of the body. Anterior setae all capillaries; posterior setae include capillaries and uncini in both rami. Scolelepis BlainvUle 1828, Lumbrkus squamatus O.F. Mller 1806; 20 species. I^stomium usually pointed, rarely blunt. Branchiae present from setiger 2, partially fiised to the notopodial (Ktstsetal lobes. Neuropodial jxjstsetal lobes notched at least in posterior setigers. Anterior setae all capillaries; posterior parapodia with uncini and capillaries in both rami. Spio Fabricius 1785, Nereis filicornis O.F. Mller 1776; 15 species. Prostomium anteriorly rounded. Branchiae present from setiger 1, at least anteriorly fused with the notopodial postsetal lobes. Notosetae all capillaries; neurosetae include capillaries and uncini. Spiophanes Grube 1860, S. kroeyeri Grube 1860, 16 species. Prostomium blunt or with frontal horns; occipital antenna present. Branchiae absent. First neuropodia with stout hooks; all other anterior setae capillaries; posterior neuropodia with uncini. Streblospio Webster 1879a, S. benedicti Webster 1879a,
2 species.
Tripolydora Woodwick 1964, T. spinosa Woodwick 1964, only species. Prostomium anteriorly rounded. Branchiae present fiom setiger 2. Fifth setiger modified, with a few modified setae only. Notosetae all capillaries; neurosetae include capillaries and trifid uncini first present from setiger 9, Taxonomic Note The above survey of the genera follows in the main Foster's (1971) survey of the fauna fixsm the Gulf of Mexico, except that I have preferred to list all the taxa at the generic level, rather than use some of them as subgenera. From the number of species listed for each genus, it will be noted that I have been rather more conservative than Foster in retaining species. The family is one of the better studied, and best luiderstood families. In addition to Poster's extensive review cited above, Hannerz (1956) and Sderstrm (1920) reviewed the major taxa. The biology of the more common forms also has been rather well studied, especially the reproductive biology of members of Polydora and Boccardia has been the subject of several studies by Blake (e.g. Blake 1971), Incertae Sedis Aberranta Hartman 1965, A, enigmtica Hartman 1965; only species. Paired spioniform palps present; prostomiimi otherwise without appendages, eyes absent. Pharynx muscularized. One apodous segment present; all others with bkamous parapodia. All setae simple capillaries. Dorsal and ventral cirri and small branchiae jwesent. According to Hartman (1%5) this differs from the true spionids in that it has a muscular, rather than saclike pharynx; it further differs from the spionids in that it completely lacks hooks of any kind. This is not a larval form, in that one of the specimens described was ovigerous. The structure of the proboscis may not be a very important feature in this group as noted by Orrhage (1966). The total absence of uncmi may not be particularly surprising, considering how many spionid genera may lack such hooks in either of the two rami, or over long stretches of the body. However, it is considered best to leave the genus as a fnee-standing genus, until it can be better investigated. Invalid Genera Aonis Audouin and Milne Edwards 1833c, xt Scolelepis Aonopsis Wagner 1885, indeterminable Aricideopsis Johnson 1901, sec Ironice Bilobaria Sveshnikov 1959, larval forms Carazzia Mesnil 1896, see Pseudopolydora Chaetosphaera Haecker 1896, larval forms
Prostomium conical. A single pair of branchiae on setiger 1. Setiger 2 with a dorsal collar between the notopodia. Notosetae all capillaries; neurosetae anteriorly all capillaries, posteriorly also uncini.
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Cheimnotus Costa 1861b, indeteraiinabie {IPolydora) Colobranchus Schmarda 1861, set Scolelepis Ctenospio Sars 1867, seePrionospio Diplotis Montagu 1815, see Polydora Dipolydora Verrill 1881, see Polydora Euspio Mclntosh 1915, seeSpio Hekaterobranchus Buchanan 1890, see Streblospio Heterospio Czemiavsky 1881b, see Spio Kinbergella Mclntosh 1909, see Prionospio Leipoceras Mbius 1874, see Polydora Leucodora Johnston 1838, see Polydora Mandarte Kinberg 1866b, indeterminable Neopygospio Berkeley and Berkeley 1954, see Pseudopolydora Nerine Johnston 1838, see Scolelepis Nerinides Mesnil 1896, see Pseudomalacoceros Nerinopsis Ehlers 1912, larval form Paranerine Czemiavsky 1881, seeAonides Perialia Kinberg 1866, questionably Boccarrfia Prospio Mesnil 1896, hypothetical Protopolydora Czemiavsky 1881b, see Polydora Pseudokucodore Czemiavsky 1881b, see Polydora Pseudonerine Czemiavsky 1881b, indetemiinable Pseudonerine Augener 1926, see Scolelepis Pteriptyches Grube 1872a, see Prionospio Pygophyllum Schmarda 1861, indeterminable Scolecolepis Malmgren 1867, mis-spelling of Scolelepis Spione rsted 1844, in Quatrefages 1865, indeterminable Spionereis Sars 1853, NOMEN NUDUM Spionides Webster and Benedict 1887, see Laonice Uncinia Quatrefages 1865, see Scolelepis 6. (A), Family MAGELONIDAE, Magelona sp., off Santa Barbara, California, diagram of dorsal view, 25x; (B), ventral view of the above, 25x.
RGURE
FAMILY MAGELONIDAE CUNNINGHAM AND RAMAGE
1888
Spioniforms with long, slender bodies, separated into two regions. Prostomium^flattened and anteriorly ovate or truncate, without appendages. Palps at the junction of the pro- and peristomium on the ventral side. Setae include capillaries and hooded bi- or multidentate hooks. Magelonids currently are assigned to a single genus, Magelona Mller 1858, with type species M. papillicornis Mller 1858 and a total of about 35 described species. Dr. Meredith L. Jones of the Smithsonian Institution is revising the family. It is anticipated that more taxa may be recognized when this revision is completed. Magelonids are common in sandy bottoms; they are rather characteristic with the flattened, shovel-shaped prostomium often much wider than the rest of the animal, which tends to be rather threadlike with very long segments. Magelonids build only very flimsy tube-structures and tend to move through the sediment.
Invalid Genera Maea Johnston 1865, sec Magelona Papillaria Sveshnikov 1959, larval fomi Rhynophylla Carrington 1865, sen Magelona
FAMILY TROCHOCHAETIDAE PETTIBONE
1963
Body flattened cylindrical, ftostomium a flattened ridge with palps present at the postectal comers. First parapodia directed forwards along the sides of the prostomium. Setae capillaries, acicular falcate spines, fringed setae and straight spines. The family is recognized for a single genus, Trochochaeta Levinsen 1884 with type species Disoma multisetosum rsted 1844 and a total of nine recognized species. Pettibone (1963) correctly pointed out that the
1976
27
name Disoma, originally applied to these animals by rsted (1844), was used ten years earlier by Ehrenberg for a protozoan. Trochochaetids are long, slender non-tubicolous spioniforms mainly reported from shallow water in soft substrates. They appear to be nowhere truly numerous, but frequently are reported from areas where quantitative benthic investigations have been undertaken. Invalid Genera Cherusca Mller 1858, indeterminable Disoma rsted 1844, see Trochochaeta Disomides Chamberlin 1919c, see Trochochaeta Nevaya Mclntosh 1911, see Trochochaeta Pilearia Sveshnikov 1963, larval forms Thaumastoma Webster and Benedict 1884, see Trochochaeta The family Kalaminochaetidae Nolte, 1941 was described for two genera, Kalaminochaeta and Kaluntmaria both based on pelagic larval forms, possibly of a Trochochaeta, but not identifiable. The family and both genera are here considered invalid.
FAMILY POECILOCHAETIDAE HANNERZ
rami. Setae include capillaries, pectinate, plumose and acicular setae. Dorsal and ventral cirri spindle or bottleshaped .
1956
Spioniforms with long, slender bodies. Prostomium small with either a frontal or a median antenna. Palps present at the postectal comers. First segment with or without setae, with one or two pairs of tentacular cirri. Parapodia biramous, with lateral sense-organs between
FIGURE 7. Family POECILOCHAETIDAE, Poecilockaetus johnsoni, off Loon Point, Santa Barbara, California, 35 m, 25x.
Key to Genera Antenna frontal, first segment with series of long setae Antenna median, first segment asetigerous Poecilochaetus Elicodasia
Generic Definitions Elicodasia Laubier and Ramos 1973, E. mirabilis Laubier and Ramos 1973; only species. Poecilochaetids with a median antenna, without nuchal organ. First segment is asetigerous and first setiger lacks dorsal and ventral cirri. Neuropodial acicular spines are present in most setigers from segment 4. Poecilochaetus Claparde 1875, P. fulgoris Claparde in Ehlers 1875; 12 species. Poecilochaetids with a frontal antenna, nuchal organs present. First segment with long setae forming a cephalic cage and second setiger similar to all following segments. Neuropodial acicular spines present in a few anterior segments only.
Taxonomic Notes Setae are present from the second segment in Elicodasia and from the first in Poecilochaetus.
FAMILY HETEROSPIONIDAE HARTMAN
1963a
Body with short thoracic and greatly prolonged abdominal setigers. Prostomium blunt without appendages; palps attached at the pwstectal comers. Long, filiform branchiae on the thoracic setigers. Setae in nearly complete cinctures around the abdomen; most simple capillaries, but some neurosetae are thickened and gently falcate. The family is known for a single genus, Heterospio Ehlers 1874 with type species H. longissima Ehlers 1874 and for a total of four species.
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Heterospionids have been reported from widely scattered areas, mainly in deep shelf and slope sediments; the family is poorly known and the several species may be considerably more common than their scattered records indicate. They are rather fragile and non-descript and thus easily overlooked if quantitative studies are not undertaken. Invalid Genus Longosoma Hartman 1944c, see Heterospio Suborder Chaetopteriformia Short palps on prostomium; pharynx a simple, noneversible tube, Uncini present.
FAMILY CHAETOPTERIDAE MALMGREN
1867
Body with two or three distinct body-regions. Peristomium may have one or two pairs of tentacular cirri; palps of varying lengths always present. Anterior region with uniramous parapodia, median and posterior regions with biramous parapodia. Setae include capillary, limbate setae and modified spines in setiger 4. Pectiniform uncini present in posterior setigers.
FIGURE 8. (A), Family CHAETOPTERIDAE, Chaetopterus variopedatus, modified from various sources, natural size; (B), Family CIRRATULIDAE, Tharyx moniloceras. after Hartman, 1969, 80x.
Key to Genera la. lb. 2a. 2b. 3a. 3b. Median notopodia bilobed Median notopodia never bilobed, may be fused A pair of small tentacular cirri present at the base of the large palps Tentacular cirri absent, palps large Some median notopodia fused to form dorsal fans, palps very short Notopodia never fused; palps large 2 3 Phyllochaetopterus Spiochaetopterus Chaetopterus Mesochaetopterus
(la) (la) (lb) (lb)
Generic Definitions Chaetopterus Cuvier 1827, Tricoelia variopedatus Renier 1804; only species. Body divided in three distinct regions; anterior region with uniramous parapodia with lancet-shaped setae, except setiger 4, which has stout spines. Median region with biramous parapodia; some posterior notopodia fused to form dorsal fans, all notopodia of median region asetigerous; neuropodia with pectinate uncini; posterior region with long, pointed notopodia with a few contained acicula, but no setae; neuropodia with pectiniform uncini. Mesochaetopterus Potts 1914, M. taylori Potts 1914; 11 species. Body with three regions; antennae absent. Median notopodia all simple, asetigerous; median neuropodia
with uncini; posterior region with pointed notopodia with internal acicula; neuropodia with uncini. Phyllochaetopterus Grube 1863, P. gracilis Grube 1863; 17 species. Body with three distinct regions; antennae present. Median notopodia foliaceous, bilobed and asetigerous, neuropodia with pectinate uncini; posterior region with pointed notopodia supported by internal acicula; neuropodia with pectinate uncini. Spiochaetopterus Sars 1853, S. typicus Sars 1856; 12 species. Body with three distinct body-regions; antennae absent. Median notopodia bilobed and foliaceous; neuropodia with pectinate uncini; posterior region with pointed notopodia, supported by internal acicula, neuropodia with pectinate uncini.
1976 Invalid Genera
29
Leptochaetopterus Berkeley 1927, see Spiochaetoplerus Mesotrocka Leuckart and Pagenstecher 1855, see Phyllochaetopterus Ranzania Claparde 1870b, see Mesockaelopterus Ranzanides Chamberlin 1919c, see Mesochaetopterus Telepsavus Costa 1861a, see Spiochaetopterus Tricoelia Renier 1804, see Chaetopterus
Suborder Cirratuliformia One pair of grooved palps, or a group of grooved palps on one or several post-peristomial segments. Prostomium without appendages, eversible pharynx a thick ventral pad.
FAMILY CIRRATULIDAE CARUS
1863
FIGURE 9. (A), Family HETEROSPIONIDAE, Heterospio catalinensis, modified from Hartman, 1944c, I2x; (B), Family ACROCIRRIDAE, Acrocirris fromifilis, after Banse, 1969, about 5x; (c), Acrocirris crassifiUs, lateral view, after Banse, 1969, about 12x,
Body cylindrical. Prostomium conical or blunt; peristomium fused with at least two segments. Parapodia reduced; slender filiform or clvate branchiae present on at least some setigers. All setae simple including capillaries and curved or excavate hooks.
Key to Genera 1 a, lb. 2a (la). 2b (la). 3a (2b). 3b (2b). 4a (3b). 4b (3b). 5a (lb). 5b (lb). 6a (5b). 6b (5b). 7a (6b). 7b (6b). 8a (7b). 8b (7b). A pair of long grooved palps attached on the anterior dorsum 2 Two groups of grooved tentacular cirri present 5 All setae slender, distally pointed Tharyx At least some setae either curved hooks or spines 3 Acicular setae distally excavate; body usually dark green or brown Dodecaceria Acicular setae not excavate, body usually light colored 4 Acicular spines in posterior segments distally entire Chaetozone Acicular spines in posterior segments distally bi- or multifid Caulleriella All setae acicular, falcate spines Pseudocirratulus At least some capillary setae present 6 Anterior region without long tentacular structures Raricirrus One or more anterior segments with groups of long tentacular cirri or branchiae 7 Dorsal tentacular cirri first present posterior to the anteriormost branchiae Cirriformia Dorsal tentacular cirri first present from the same segment as the anteriormost branchiae 8 Tentacular cirri on one segment only Cirratulus Tentacular cirri on two or more segments Timareie Generic Definitions Caulleriella Chamberlin 1919c, Cirratulus bioculatus Keferstein 1862; 16 species. Paired palps inserted dorsally anterior to, or at the first setiger. Setae include capillaries and distally bidentate or multidentate curved spines. Chaetozone Malmgren 1867, C. setosa Malmgren 1867; 19 species. Paired palps inserted dorsally anterior to, or at the first setiger. Setae include capillaries and distally entire, curved spines. Cirratulus Lamarck 1801, Lumbricus cirratus O.F. Mller 1776, 25 species. Cirratulids without palps; tentacular cirri present on one segment only; anteriormost branchiae present from the same segment. Setae include capillaries and acicular spines that are usually entire distally.
30
NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY Taxonomic Notes
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Cirriformia Hartman 1936a, Terebella tentaculata Montagu 1808; 26 species. Cirratulids without palps; tentacular cirri present on one segment only; anteriormost branchiae present anterior to the tentacular cirri. Setae include capillaries and curved, distally entire spines. Dodecaceria rsted 1843b, D. concharum rsted 1843b; 16 species. Palps present; branchiae present on a few segments; filiform or clvate. Anterior setigers with capillary setae; median and posterior ones also with excavate stout spines. Posterior part of body often flattened and expanded. Pseudocirratulus Augener 1922, P. kingstonensis Augener 1922; only species. Palps absent, cirri and branchiae present on most segments; two achaetous anterior segments present. Setae all gently curved, entire spines or hooks. Raricirrus Hartman 1961, R. maculatus Hartman 1961; only species. Body anteriorly narrow, then abruptly broad at seliger 9 and tapering from there to the posterior end. Branchiae present on some anterior segments; setiger 13 with a pair of papillae. Setae include capillaries in the notopodia and neuropodia and curved hooks with dentate cutting edges. Tharyx Webster and Benedict 1887, T. acutus Webster and Benedict 1887; 22 species. Palps present anterior to or at the first stiger. All setae slender, mostly capillary and smooth, but sometimes with serrated cutting edges. Timarete Kinberg 1866b, Cirratulus anchylochaetus Schmarda 1861; 8 species. Palps absent; tentacular cirri present on at least two anterior setigers; branchiae present from the same segment as the first tentacular cirri. Setae include notopodial capillaries and anteriorly neuropodial capillaries; posterior neuroopodia with gently curved spines.
It appears likely that the generic division of the palpate cirratulids will change with future revisions. The TAaryJc-complex is presently rather confused, and additional characters will have to be included to clarify the relations between the several species described in this genus.
Invalid Genera Ambo Chamberiin 1918, see Timarete Archidice Kinberg 1866b, see Cirratulus Audouinia Quatrefages 1865, see Cirriformia Cirratulispio Mclntosh 1911, questionably Cirratulus or Chaetozone Cirrhineris Blainville 1818, indeterminable Heterocirrus Grube 1855, $&e Dodecaceria Heterocirrus Saint-Joseph 1894, see Caulleriella and in part Tharyx Labranda Kinberg 1866b, see Cirriformia Mesocirrineris Czerniavsky 1881b, i&e, Caulleriella Monticellirux Laubier 1961a, see Tharyx Naraganseta Leidy 1855, see Dodecaceria Pentacirrus Wesenberg-Lund 1958, see Timarete Promenia Kinberg 1866b, see Cirratulus
FAMILY ACROCIRRIDAE BANSE
1969
Body either slender and elongate, or short and maggot-shaped. Prostomium blunt, paired palps usually present. Several segments usually crowded near the anterior end; these segments usually asetigerous. Parapodia biramous with small parapodia; notosetae segmented and spinose; neurosetae compound hooded falcigers. The acrocirrids recently were recognized as a separate family, and even more recently, the genus Flabelligella known from several deep-water locations was transferred to this family (Orensanz 1974b).
Key to Genera la. 1 b. 2a (lb). 2b (lb). Branchiae absent Branchiae present Palpal bases well separated; epithelium densely papillated Palpal bases nearly abutting, epithelium nearly smooth Flabelligella 2 Macrochaeta Acrocirrus
Generic Definitions Acrocirrus Grube 1872b, Heterocirrus fi-ontifilis Grube I860; 8 species. Acrocirrids with palps originating close together. Four pairs of gills; a papilla on third segment below
the gill. Thorax of 12 segments; a row of papillae ventral to the neurosetae in all setigers. Flabelligella Hartman 1965, F. papillata Hartman 1965; 5 species. Acrocirrids without branchiae and palps; separation
1976
i]
between thorax and abdomen indistinct. Body-surface densely papillated. Anterior segments uniramous in some species. Macrochaeta Grube 1850, Nais clavicomis Sars 1835; 6 species. Acrocirrids with bases of palps separated by at least the width of the palps. Usually four pairs of branchiae. Number of thoracal segments variable; epidermis usually densely papillated. All parapodia biramous. Invalid Genus Ledon Webster and Benedict 1887, ^t& Macrochaeta ORDER CAPITELLIDA Prostomium without appendages; palps absent. One or two anterior asetigerous segments. Parapodia biramous; neuropodia long transverse welts in at least some setigers. All setae simple, including capillaries and rostrate uncini.
FAMILY CAPITELLIDAE GRUBE
difficult. However, genera and species cannot be identified without this information. 1 have found it useful to rotate the specimen while counting the segments, trying to observe the setae as the light catches them. The rostrate uncini look truncate in this sort of observation, and the capillaries will retain their tapered appearance,
1862
Body cylindrical, usually slender. Prostomium short and truncate, without appendages. Thorax and abdomen defined on internal structures and parapodial development; generally, thorax with capillary setae and abdomen with rostrate hooded hooks. Capitellids are among the most commonly encountered polychaetes; some forms, such as Capitella capitala are considered pollution indicators, in that they are capable of invading areas where disastrous defaunation of natural or man-made causes has taken place (Grassle and Grassle 1974). The anterior thorax of most capitellids is usually strongly areolated and they are usually reddish pink or brown. In general appearance, they are perhaps the most earthwormlike of the common marine polychaetes. The characteristic rostrate uncini are small, making the determination of the first occurrence of these setae
10. (A), Faraiiy CAPlTEhLlDAB, Noiomasms lenuis. after Hartman, 1947, 7x; (b), hooded hook of the above, )i50x; (C), Family ARENICOLIDAE,.4rert(fo/fl, near marina, modified from several sources, about natural sire.
FIGURE
Key to Genera la. lb. 2a (la). 2b (la). 3a (2b). 3b (2b). 4a (lb). 4b (lb). 5a (4a). 5b (4a). Three first setigers with capillary setae only At least four setigers with capillary setae only Some thoracic segments with flanged hooks; twelve thoracic segments present Flanged hooks absent; either ten or thirteen thoracic segments present Setigers 7-8 with genital spines; ten thoracic segments Genital spines absent; 13 thoracic segments Four first setigers with capillary setae only At least five anterior setigers with capillary setae Genital spines present Genital spines absent 2 4 Peresiella 3 Capitomastus 9 Heteromastides 5 8 6 7
32 6a (5a). 6b (5a). 7a (5b). 7b (5b). 8a (4b). 8b (4b). 9a (8a). 9b (8a). 10a (8b). 10b (8b). lia (10a). I lb (10a). 12a (11 b). 12b (lib). I3a (10b). 13b (10b). 14a (13a). 14b (13a). 15a (14a). 15b (14a). 16a (14b). 16b (14b). 17a (13b). 17b (13b). 18a (17a). 18b 19a 19b 20a 20b 21a ( 17a). (17b). (17b). ( 19a). (19a). (20b).
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21b (20b). 22a (21b). 22b (21b). 23a 23b 24a 24b 25a (19b). (19b). (23a). (23a). (24a).
25b (24a). 26a (24b). 26b (24b). 27a (23b). 27b (23b). 28a (27a). 28b (27a). 29a (28b). 29b (28b). 30a (27b). 30b (27b).
Genital spines in setigers 7-8; ten thoracic segments present Capitomastus 6 Genital spines in setigers 8-9; nine thoracic segments present Capitella (part) Thorax with 11 segments Mediomastus Thorax with 12 segments Parheteromastus Five setigers with capillary setae 9 At least six anterior setigers with capillary setae only 10 First setiger with both noto- and neuropodia; branchiae present Heteromastus First setiger with notopodia only; branchiae absent Parheteromastides Six anterior setigers with capillary setae only 11 At least seven anterior setigers with capillary setae 13 Genital spines in setigers 8-9 Capitellides Genital spines absent 2 Thorax with 11 segments Neomediomastus Thorax with 12 segments Barantolla Seven anterior setigers with capillary setae only 14 At least nine anterior setigers with capillary setae only 17 Genital spines present 15 Genital spines absent 16 Notopodial cirriform branchiae on some abdominal segments Branchiocapitella Branchiae absent Capitella Thorax with eight segments; all thoracic setigers with capillary setae only LeiocapitelUdes Thorax with 12 segments; posterior thoracic setigers with hooks Neoheteromastus Nine anterior setigers with capillary setae only 18 Ten or more anterior setigers with capillary setae 19 One anterior asetigerous segment present; notopodial acicula absent; thorax with ten segments Pseudoleiocapitella Anterior asetigerous segment absent; notopodia with acicula; thorax with nine segments . . , Pulliella Ten first setigers with capillary setae only 20 At least 11 anterior setigers with capillary setae 23 First setiger with both noto- and neuropodia Decamastus First setiger with notopodium only 21 All thoracic setigers with capillary setae; all abdominal setigers with hooks only; mixed segments absent Capitellethus At least one segment with mixed hooks and capillary setae 22 Two first abdominal segments with mixed hooks and capillary setae; thorax with eleven segments Neonotomastus Last thoracic segment with both hooks and capillary setae; twelve thoracic segments present Paraleiocapitella Eleven anterior setigers with capillary setae only 24 Twelve or more anterior setigers with capillary setae only 27 At least some capillary setae in the first two abdominal setigers 25 All abdominal segments with hooks only 26 First setiger with neuropodia only; first two abdominal setigers with capillary setae in both rami Notodasus First setiger with both noto- and neuropodia; first two abdominal setigers with hooks in the neuropodia and mixed hooks and capillary setae in the notopodia Mastobranchus Abdominal notopodial tori nearly coalesce; notopodial hooks absent Rashgua Abdominal notopodial tori well separated; notopodial hooks present Notomastus Twelve setigers with capillary setae only 28 Thirteen or more setigers with capillary setae only 30 Anal plaque with imbedded spines present Scyphoproctus Anal end rounded or with anal cirri 29 Thirteenth setiger with both hooks and capillary setae Leiochrus No setiger with both hooks and capillary setae present Leiochrides Thirteen anterior setigers with capillary setae only 31 At least 15 anterior setigers with capillary setae 35
1976 31a (30a). 31b (30a). 32a (31a). 32b (31a). 33a 33b 34a 34b 35a (31b). (31b). (33b). (33b). (30b).
ii
35b (30b). 36a (35b). 36b (35b). 37a (36b). 37b (37b).
First seliger with notopodia only 32 First setiger with both noto- and neuropodia 33 Thorax with 14 or 15 segments; one segment with both capillary setae and hooks present Leiocapitella Thorax with 15 to 18 segments; three or four segments with both capillary setae and hooks present Pseudocapitella Thorax with 19 segments; branchiae neuropodial Capitobranchus Thorax with maximally 15 segments; branchiae, if present, notopodial 34 All notopodial setae capillary Protomastobranchus Abdominal notopodial setae hooks Dasybranchus At least 16 setigers with capillary setae only; additional four segments have both setae and hooks in the notopodia Eunotomastus Thoracic count otherwise; maximally two transitional segments present 36 First setiger with both noto- and neuropodia Dasybranchetus First setiger with notopodia only 37 Twenty setigers with capillary setae Lumbricomastus Seventeen or 18 setigers with capillary setae only Anotomastus
Generic Definitions Anotomastus Hartman 1947, Eunotomastus gordioides Moore 1909; only species. Thorax with 18 or 19 segments; one anterior asetigerous segment present; first setiger with notopodia only. Up to segment 17 or 18 capillary setae only; segments 18 and 19 transitional with notopodial setae and neuropodia with mixed hooks and setae. Palmately branched branchiae in posterior segments. BarantoUa Southern 1921, B. sculpta Southern 1921; 2 species. Thorax with 12 segments; one anterior asetigerous segment present; first setiger complete; up to segment 7 with capillary setae only; segment 8 with mixed setae and hooks in notopodia and hooks in neuropodia. Branchiae may be present in posterior segments. Branchiocapitella Fauve I 1932, B. singularis Fauvel 1932; 2 species. Thorax with nine segments; asetigerous segment absent; first setiger complete. Up to segment 7, capillary setae only; segments 8-9 with genital spines. Notopodial cirriform branchiae present. Capitella Blainville 1828, Lumbricus capitatus Fabricius 1780; 15 species. Thorax with nine segments; asetigerous segment absent; first setiger complete. Either four first segments with capillary setae only; then three segments with mixed hooks and setae in both rami, then genital spines in segments 8-9, or first seven segments with capillary setae only; then segments 8-9 with genital spines. Branchiae absent. Capitellethus Chamberlin 1919c, CapitelUdes dispar Ehlers 1907; 3 species.
Thorax with 11 segments; asetigerous segment absent; first setiger with notopodia only. All thoracic setigers with capillary setae only; branchiae absent. CapitelUdes Mesnil 1897b, C. giardi Mesnil 1897b; 3 species. Thorax with nine segments; asetigerous segment absent; first setiger complete. First six setigers with capillary setae only; next three with hooks, except notopodia 8-9, which has genital spines. Branchiae absent. Capitobranchus Day 1962, C. macgregori Day 1962; only species. Thorax with 19 segments; one asetigerous segment present; first setiger complete. Up to segment 15 with capillary setae only; next four segments mixed with capillary setae in the notopodia and hooks in the neuropodia. Neuropodial branchiae present in posterior setigers. Capitomastus Eisig 1887, Capitella minima Langerhans 1881; 3 species. Thorax with ten segments; one asetigerous segment present; first setiger complete. Males: up to segment 5 with capillary setae only, the next two segments with hooks, segments 8-9 with genital spines. Females: up to segment 4 with capillary setae; the next three segments with hooks and segments 8-9 with genital spines. Dasybranchetus Monro 1931, D. fauveli Monro 1931; only species. Thorax with 16 segments; one asetigerous segment present; first setiger complete; all thoracic setigers with capillary setae. Branchiae not seen. Dasybranchus Grube 1850, Dasymallus caducus Grube 1846; 10 species.
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Thorax with 14 segments; one asetigerous segment present; first seliger complete; all thoracic setigers with capillary setae only. Simple or composite notopodial branchiae present. Decamastus Hartman 1963a, D. gracilis Hartman 1963a; 2 species. Thorax with 11 segments; one asetigerous segment present; first setiger complete; all thoracic setigers with capillary setae only; branchiae absent. Eunotomastus Mclntosh 1885, E. grbet Mclntosh 1885; only species. Genus poorly defined; approximately 20 thoracic segments present; of which 16 have capillary setae only and the next four have mixed setae and hooks in the notopodial fascicles. Heteromastides Augener 1914, H. bifidus Augener 1914; 2 species. Thorax with 13 segments; one asetigerous segment present; first setiger complete. Up to segment 4, capillary setae only; segment 5 with mixed capillary setae and hooks in both rami; from segment 6, hooks only. Branchiae absent. Heteromastus Eisig 1887, Capitella filiformis Claparde 1864; 7 species. Thorax with 12 segments; one asetigerous segment present; first setiger complete. Up to segment 6, capillary setae only; from segment 7, all setae hooks. Notopodial branchiae present. Leiocapitella Hartman 947, L. glabra Hartman 1947; 2 species. Thorax with 14 or 15 segments; one asetigerous segment present; first setiger with notopodia only. Up to segment 14 with capillary setae only; segment 15 with capillary setae in the notopodia and hooks in the neuropodia. Branchiae absent. Leiocapitellides Hartmann-Schrder 1960a, L. analis Hartmann-Schrder 1960a; only species. Thorax with eight segments; one asetigerous segment present; alt thoracic segments with capillary setae only; first abdominal with notopodial capillary setae and neuropodial hooks. Branchiae absent. Leiochrides Augener 1914, L. australis Augener 1914; 7 species. Thorax with 13 segments; one asetigerous segment present; first setiger complete. All thoracic setigers with capillary setae only. Branchiae not seen. Leiochrus Ehlers 1908, L. alutaceus Ehlers 1908; only species. Thorax with 13 or 14 segments; one asetigerous segment present; first setiger complete. Up to segment 13, capillary setae only; segment 14 with mixed setae and hooks in both rami. Branchiae absent.
Lumbricomastus Thomassin 1970, L. tulearensis Thomassin 1970; only species. Thorax with 21 segments; one asetigerous segment present; first setiger with notopodia only. All thoracic setigers with capillary setae only. Branchial processes on posterior notopodia. Mastobranchus Eisig 1887, M. trinchesii Eisig 1887; 4 species. Thorax with 12 segments; one asetigerous segment present; first setiger complete. All thoracic setigers with capillary setae only; two first abdominal segments with mixed capillary setae and hooks in the notopodia and hooks in the neuropodia. Notopodial branchiae present. Mediomastus Hartman 1944b, M. californiensis Hartman 1944b, 7 species. Thorax with 11 segments; one asetigerous segment present; first setiger complete. Up to segment 5 with capillary setae only; thereafter, all hooks. Branchiae absent. Neoheteromastus Hartman 1960, N. lineus Hartman 1960; only species. Thorax with 12 segments; one asetigerous segment present; first setiger with notopodia only. Up to segment 8 with capillary setae only; segment 9 with capillary setae in the notopodia and hooks in the neuropodia. Branchiae not seen. Neomediomastus Hartman 1969, Mediomastus glabrus Hartman 1960; only species. Thorax with 11 segments; one asetigerous segment present; first setiger complete. Up to segment 7 with capillary setae only, then all setigers with hooks. Small notopodial branchial processes present in far posterior setigers. Neonoiomastus Fauchald 1972, N. glabrus Fauchald 1972; only species. Thorax with 11 segments; one asetigerous segment present; first setiger with notopodia only. All thoracic setigers with capillary setae only; first abdominal notopodia with capillary setae, second with mixed hooks and capillary setae. First abdominal neuropodia with mixed hooks and capillary setae; second with hooks only. Branchiae not seen. Notodasus Fauchald 1972, N. magnus Fauchald 1972; 2 species. Thorax with 12 segments; one asetigerous segment present; first setiger with neuropodia only. All thoracic setigers and the first two abdominal setigers with capillary setae only. Branchiae not seen. Notomastus Sars 1850, N. latericeus Sars 1850; 34 species. Thorax with 12 segments; one asetigerous segment
]976
35
present; first se tiger complete. All thoracic setigers with capillary setae only. Branchiae may be present. Paraleiocapitella Thomassin 1970, P. mossambica Thomassin 1970; only species. Thorax with 12 segments; one asetigerous segment present; first setiger with notopodia only. Up to segment II with capillary setae only; segment 12 with notopodial capillary setae and neuropodial hooks. Branchiae absent. Parheteromastides Hartmann-Schrder 1962a, P. multiocuiatus Hartmann-Schrder 1962a; only species. Thorax with 11 segments; one asetigerous segment present; first setiger with notopodia only. Up to segment 6, capillary setae only; segment 7 with both capillary setae and hooks and neuropodia with hooks only. Branchiae absent. Parheteromastus Monro 1937a, P. tenuis Monto 1937a; only species. Thorax with 12 segments; one asetigerous segment present. Up to segment 5, capillary setae only; all other segments with hooks only. Branchiae absent. Peresiella Harmelin 1968. P. clymenoides Harmelin 1968; 2 species. Thorax with 12 segments; one asetigerous segment present; first setiger with notopodia only. Up to segment 4, capillary setae only; the remainder of thorax with modified, flanged hooks in at least some segments, capillary setae and normal hooks in the others. Protomastobranchus Gallardo 1968, P. huloti Gallardo 1968; only species. Thorax with 13 or 14 segments; one asetigerous segment present; first setiger complete. All notopodia with limbate capillary setae only; up to segment 13 or 14, capillary setae in neuropodia also; from there on, hooks. Pseudocapitella Fauvel 1913, P. incerta Fauvel 1913; 2 species. Thorax with 15 to 18 segments; one asetigerous segment present; first setiger with notopodia only. Up to segment 15, capillary setae only; next three with notopodial capillary setae and neuropodial hooks. Branchiae absent. Pseudoleiocapitella Harmelin 1964, P. fauveli Harmelin 1964; only species. Thorax with ten segments; one asetigerous segment present; first setiger complete. Up to segment 10, capillary setae only; first two abdominal segments with notopodial capillary setae and neuropodial hooks. Branchiae absent. Pulliella Fauvel 1929, P. armata Fauvel 1929; 2 species. Thorax with nine segments; all with capillary setae.
First setiger complete. Branchiae absent; notopodial acicula present. Rashgua Wesenberg-Lund 1949, P. rubrocinaa Wesenberg-Lund 1949; only species. Thorax with 12 segments; one asetigerous segment present; first setiger complete; all thoracic setigers with capillary setae. Dorsal abdominal tori nearly coalesce, lack notopodial hooks. Simple branchiae present. Scyphoproctus Gravier 1904, 1904; 7 species. Thorax with 13 segments; present; first setiger complete. capillary setae only. Expanded imbedded spines present. S. djiboutensis Gravier one asetigerous segment All thoracic setigers with anal plaque with acicular
Taxonomic Notes The generic sub-division of the capitellids is unsatisfactory; it is based on the number of thoracic segments, which may be difficult to see, and on the distribution of the capillary and hooked setae, which is easily enough seen, but poorly understood in terms of variability. The key to genera follows the traditional system, but has been based on the number of segments with capillary setae, rather than on the number of thoracic segments, since the former is the more easily observed character. A review only, such as the current one, cannot solve the problem of the number of valid capitelHd genera, which must be based on the variability of all observable characters in a large amount of material. The genus Capitita Hartman 1947, is considered here a synonym of Mediomastus Hartman 1944b, as first suggested by Hartmann-Schrder (1962a). The genus Bucherta Rullier (1965a) is considered as the posterior end of a capitellid, probably of the genus Dasybranchus as first suggested by Gallardo (1968). As stated above, the generic identification of capitellids is at best difficult. To make a review somewhat easier, a table has been constructed to indicate in a different manner the relationships between the genera. It is of the utmost importance that the two concepts, segments and setigers, be kept apart, since both are used in key features of the capitellids.
Invalid Genera Ancistria Quatrefages 1865, ise Heteromastus Arenia Quartrefages 1865, se& Noiomasius Areniella Verrill 1874, questionably//eromajfujBranchoscolex Schmarda 1861, s^s Dasybranchus Bucherta Rullier 1965a, see Dasybranchus
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Capitellides Ehlers 1907, see Capitellethus Dasymallus Grube 1846, see Dasybranchus Ditrocha Sveshnikov 1959, larval fonns Eisigella Gravier 1901, see Notomastus
Isomastus Gravier 191 la, see Capitella Lumbriconais rsted 1842, in Grube 1850, stt Capitella Sandanis Kinberg 1867b, set Notomastus Valla Johnston 1865, see Capitella
TABLE 2 Genera of Capitellidae. Airanged in order of increasing number of thoracic segments present.
Column 1: No. of thoracic segments present Column 2: Presence or absence of an anterior asetigerous segment Column 3; No. of thoracic setigers Column 4: Essence of genital spines
Genus Leiocapitellides Puiiiella Capitella (1) Capitellides Capitella (2) Branchiocapitella Capitomastus 9 Capitomastus 6 Pseudoleiocapitella Mediomastus Parheteromastides Neomediomastus Capitellethus Decamastus Neonotomastus Peresiella Parheteromastus Heteromastus BarantoUa Neoheteromastus Paraleiocapitella Notomastus Rashgua Mastobranchus Notodasus Heteromastides Leiochrides Scyphoproctus Leiocfirus Protomastobranchus Dasybranchus Leiocapitella Pseudocapitetla Dasybrancketus Anotomastus Capilobranchus Lumbricomastus 1 8 9 9 9 9 9 10 10 10 11 11 11 11 11 11 12 12 12 12 12 12 12 12 12 12 13 13 13 13-14 13-14 14 14-15 15-18 16 18-19 19 21 2 3 7 9 9 9 9 9 9 9 9 10 10 10 10 10 10 11 11 11 11 11 11 11 11 11 11 12 12 12 12-13 13-14 13 13 14-17 15 17-18 18 20 4 5 7 9 4 6 7 7 3 4 9 4 5 6 10 10 10 3 4 5 6
Column 5: No. of setigers with capillary setae Column 6: No. of setigers with mixed capillaries and rostrate uncini Column 7: First segment complete or incomplete Column 8: EYesence of branchiae and special kinds of setae.
6 7 compl compl compl compl compl compl compl compl compl compl notop. compl notop. compl notop. notop. compl compl compl notop. notop. compl notop. compl compl neuro compl compl compl compl compl compl notop. notop. compl notop. compl notop. no branchiae no branchiae notacicula prs no branchiae no branchiae no branchiae branchiae no branchiae no branchiae no branchiae no branchiae no branchiae small knobs no branchiae ?no branchiae ?no branchiae Hanged setae no branchiae branchiae branchiae Tbranchiae no branchiae branchiae no notouncini branchiae branchiae ?branchiae no branchiae 7branchiae anal plaque no te-anchiae no notouncini branchiae no la-anchiae no branchiae ?branchiae branchiae neurop. branchiae branchiae 8
+ _ + + + + + + +
_
8-9 8-9 8-9 8-9 8-9 8-9
_
3
_ 2
_ _ _ _ _ _ _ _ -
_
1
_ 2 var.
+ +
+
+
+
_ 1 1 1
+
+ +
7
10 11 11 11 13 3 12 12 12 13-14 13 13 14 15 17-18 14 20
+ + +
+ +
_
2
1 1
+
+
+ + + + + + + +
1 3
1 4
The genus Eunotomastus is too incompletely known to be included in the table.
1976
FAMILY ARENICOLIDAE JOHNSTON
37
1835
Body cylindrical, separated into two or three distinct regions. Prostomium without appendages. Notopodia bluntly truncate, neuropodia elongated tori. Notosetae capillary or limbate, neurosetae rostrate hooks. Branchiae present on some setigers in a median or posterior region.
With the exception of Branchiomaldane, the arenicolids are not easily confused with any other polychaetes. The very thick, strongly areolated epidermis in most forms, the distinct branchial region with their strongly tufted branchiae and the habitat, makes it easy to recognize the "sand-worms" from all over the world. The family has been the object of very intensive studies by Wells and his students. Wells (1%2) and other papers established the major classificatory criteria.
Key to Genera la. 1 b. 2a (lb). 2b (lb). 3a (2b). 3b (2b). Body slender, branchiae first present from setiger 18, or later, as thick filaments arranged with maximally two or three in a tuft Branchiomaldane Body thick, branchiae first present from a more anterior setiger as thick tufts of very fine filaments . 2 Asetose caudal end absent Arenicolides Asetose caudal end present 3 Neuropodia of branchial segment approach midventrally; a single pair of oesophageal sacs present Arenicola Neuropodia of branchial segments well separated; more than one pair of oesophageal sacs present Abarenicola
Generic Definitions Abarenicola Wells 1959, Arenicola claparedii Levinsen 1884; 16 species and subspecies. Three body-regions, including a prebranchial and branchial region as well as an asetose caudal end. Branchiae from setiger 7. Neuropodia widely separated in the branchial region. More than one pair of oesophageal sacs. Arenicola Lamarck 1801, Lumbricus marinus Linnaeus 1758; 9 species and subspecies. Three body-regions, including a prebranchial and branchial region as well as an asetose caudal end. Branchiae fix)m setiger 7. Neuropodia approach midventrally in branchial segments. One pair of oesophageal sacs present. Arenicolides Mesnil 1898, Arenicola ecaudata Johnston 1835; 2 species. Two body-regions, including a prebranchial and a branchial region. Branchiae from setiger 12-17 to the posterior end. All neuropodia approach midventrally. Branchiomaldane Langerhans 1881, . vitwenti Langerhans 1881; only species. Two body-regions, including a prebranchial and branchial region. Branchiae first present fix)m setiger 18 or behind; as thick, sessile filaments, usually no more than three filaments in a group. All neuropodia widely separated.
Invalid Genera Chorizobranchus Quatrefages 1865, sec Arenicola Clymenides Claparde 1863, see Arenicolides Protocapitella Berkeley and Berkeley 1932, see Branchiomaldane Pteroscolex Ltken 1864, &ee Arenicola Papara Sveshnikov 1959, larval forms Telethusae Savigny 1818, see Arenicola
FAMILY MALDANIDAE MALMGREN
1867
Capitelliform polychaetes with long cylindrical bodies, usually truncate at one or both ends; most species with long, cylindrical segments. Prostomium without appendages, with a pair of nuchal slits and a median cephalic keel. Notopodia short and rounded; neuropodia elongated tori. Notosetae smooth or spinose capillaries; neurosetae rostrate hooks, anterior modified spines present in several forms. The bamboo-worms are recognized easily in that for once, the popular name of the members of this family makes sense; the long segments with the parapodia at one end, give the worms the jointed appearance of slender bamboo-shoots. However, even though easily recognized to family, they are not as easily identified to genus and species. It is necessary to have complete specimens to get them even to sub-family; or at least, one must have both anterior and posterior ends from the same specimen to get them safely iden-
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tified. It is here strongly suggested that identification of fragments not be undertaken, except of course when 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 subfamilies 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 prostomiura 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 without collars; uncini in single rows. NICOMACHINAE: Anterior end without plaque; anal plaque present; uncini in single rows. MALDANINAE: Both anterior and posterior ends with plaques; anus dorsal.
EUCLYMENINAE: Anterior and posterior ends with plaques, anus terminal.
^^^-* ]
FIGURE 11. (A), Family MALDANIDAE, Axiothella rubrocincta. Tmales Bay, California, intertidal, 17x; (B), posterior end of the above, 17x.
Key to Genera la. lb. 2a (la). 2b (la). 3a (2b). 3b (2b). 4a (3b). 4b (3b). 5a (4b). 5b (4b). 6a (5b). 6b (5b). 7a (lb). 7b (Ib). 8a (7a). 8b (7a). 9a (8b). 2 7 RHODININAE Rhodine Rostrate uncini in single rows, posterior segments not collared LUMBRICLYMENINAE 3 More than 20 setigers present Praxillura Nineteen setigers present 4 Setiger 4 with a deep encircling collar Clymenopsis Setiger 4 without collar 5 Pygidium with anus at the end of a simple, upturned, flattened cone Lumbriclymenella Pygidium otherwise 6 Pygidium flattened with anus dorsal Notoproctus Pygidium conical with anus terminal Lumbriclymene Cephalic plaque absent, anal plaque present NICHOMACHINAE 8 Both cephalic and anal plaques present 10 Rostrate uncini in all setigers Micromaldane Acicular spines in first three setigers, thereafter rostrate uncini 9 Anal funnel symmetrically developed Nicomache Both cephalic and anal plaques absent At least anal plaque present Rostrate uncini in double rows, posterior segments with encircling collars
1976 9b (8b). 10a (7b). 10b (7b). 11a (10a). lib (10a). 12a (1 lb). 12b (lib). 13a (12b). 13b (12b). 14a (13b). 14b (13b). 15a (10b). 15b (10b) 16a (15b). 16b (15b). 17a (16a). 17b (16a). 18a (16b). 18b (16b). 19a (18a). 19b (18a). 20a (19b). 20b (19b). 21a (20a). 21b (20a). 22a (20b). 22b (20b). 23a (18b). 23b (18b). 24a (23a). 24b (23a). 25a (23b). 25b (23b). 26a (25b). 26b (25b). 27a (26b). 27b (26b). 28a (27b). 28b (27b). 29a (28b). 29b (28b).
THE E*OLYCHAETE WORMS
i9
Anal funnel asymmetrical, with the dorsal side reduced Petaloproctus Anus dorsal MALDANINAE 11 Anus terramal EUCLYMENINAE 15 Rostrate uncini in two or three series in most setigers Sonatsa Rostrate uncini always in single series 12 First three setigers with acicular neuropodial spines Clymaldane First setiger either without neurosetae or with rostrate uncini 13 Cephalic keel long and high, cephalic rim shallowly notched laterally Maldane Cephalic keel short and low, cephalic rim deeply incised laterally 14 Numerous branchial filaments on median setigers Branchioasychis Branchial filaments absent Asychis Series of vascular caeca cover the surface of the two last setigers Johnstonia Vascular caeca absent 16 Anal plaque marginally smooth or gently crenulated 17 Anal plaque bordered by distinct anal cirri 18 First setiger with notosetae only; anal plaque gently crenulated Abyssoclymene First setiger with noto- and neurosetae; anal plaque smooth Microclymene All anal cirri similar in length 19 Midventral annal cirrus (rarely two) distinctly longer than all other anal cirri 23 More than 30 setigers present Macroclymene Nineteen or 20 setigers present 20 Acicular spines in the first neuropodia 21 First neurosetae rostrate uncini 22 Setiger 4 with a deep encircling collar Clymenella Setiger 4 without a collar Isocirrus First setiger with notosetae only; anal plaque with numerous cirri Maldanella First setiger with both noto- and neurosetae; anal plaque with a few long cirri only Clymenura More than 30 setigers present 24 Eighteen to 20 setigers present 25 Setiger 4 with a deep encircling collar Macroclymenelia Setiger 4 without a collar Gravierella Anterior neuropodia with rostrate uncini Axiothella Anterior neuropodia with acicular spines 26 Anal plaque with two large ventral and several shorter lateral and dorsal anal cirri Proclymene A single large ventral cirrus and several shorter lateral and dorsal anal cirri present 27 Nuchal slits short, straight and diverging anterioriy Pseudoclymene Nuchal slits long, straight and parallel 28 Anal cone projecting well beyond the rim of the anal plaque Praxillella Anal cone low and not projecting beyond the rim of the anal plaque 29 Apart from the long ventral anal cirrus, all other anal cirri similar in length Euclymene Anal cirri of varying lengths Heteroclymene
'
Generic Definitions
Abyssoclymene Hartman 1967, A. annularis Hartman 1967; only species. EUCLYMENINAE. Nuchal slits straight. Nineteen setigers, first with notosetae only; anal plaque nearly smooth, gently crenulated along margin. Asychis Kinberg 1867b, A. atlanticus Kinberg 1867b; 28 species. MALDANINAE. Cephalic rim with deep lateral incisions; cephalic keel short and low. Neurosetae absent in first setiger. Branchiae absent.
Axiothella Verrill 1900, Axiothea catenata Malmgren 1865; 12 species. EUCLYMENINAE. Cephalic rim incised or entire; 18-20 setigers, acicular spines absent. Anal plaque with long midventral and shorter lateral and dorsal anal
Branchioasychis Monro 1939c, B. colmani Monro 1939c; 3 species. MALDANINAE. Cephalic rim deeply incised laterally; cephalic keel short and low. Neurosetae absent in first setiger. Numerous gill filaments on median setigers.
40
NATURAL H/STORY MUSEUM OF LOS ANGELES COUNTY
Science Series 28
Clymaldane Mesnil and Fauvel 1939, C. sibogae Mesnil and Fauvel 1939; 2 species. MALDANINAE. Cephalic rira shallowly notched. Keel short and very low. Acicular spines present in anterior neuropodia. Branchiae absent. Clymenella Verrill 1873b, Clymene torquatus Leidy 1855; 7 species. EUCLYMENINAE. Eighteen to 20 setigers; nuchal slits long and straight. Setiger 4 with deep encircling anterior collar; anterior neuropodia with acicular spines or strongly reduced uncini. Anal plaque with all cirri evenly long, Clymenopsis Verrill 1900, Clymene cingulata Ehlers 1887; 2 species. LUMBRICLYMENINAE, Nineteen setigers; setiger 4 with deep encircling collar; nuchal slits strongly angled. First three neuropodia with spines. Clymenura Verrill 1900, Clymene cirrata Ehlers 1887; 11 species. EUCLYMENINAE. About 20 setigers; with large triangular glandular field on setiger 8. Nuchal slits long and straight. Acicular spines absent. Anal plaque a flattened disc with few long anal cirri; anal cone strongly projecting. Euclymene Verrill 1900, Clymene oerstedii Claparde 1863; 27 species. EUCLYMENINAE. Eighteen to 20 setigers; nuchal slits long and straight. Acicular spines present. Anal plaque with one long median and numerous evenly long anal cirri; anal cone low. Gravierella Fauvel 1919, G. multiannulata Fauvel 1919; only species. EUCLYMENINAE. Numerous setigers, far posterior ones campanulate. Nuchal slits long and straight. Acicidar spines absent. Anal plaque with the anal cirri increasing in length from the dorsal side towards the long midventral cirrus. Heteroclymene Arwidsson 1907, H. robusta Arwidsson 1907; 3 species. EUCLYMENINAE. Nineteen setigers; nuchal slits long and straight. Acicular spines in first neuropodia. Anal plaque with one long midventral and numerous shorter anal cirri of varying lengths. Anal cone low. Isocirrus Arwidsson 1907, Clymene planiceps Sars 1872; 5 species. EUCLYMENINAE. About 20 setigers; nuchal slits long and straight. Acicular spines in anterior neuropodia. Anal plaque with all cirri of the same length. Johnstonia Quatrefages 1850b, J. clymenoides Quatrefages 1850b; 2 species. EUCLYMENINAE. Twenty-two setigers; nuchal slits long and straight. Acicular spines in the first neuro-
podia. Anal plaque with one long midventral and several shorter anal cirri. Two last setigers covered with series of vascular caeca. Lumbriclymene Sars 1872, L. cylindricauda Sars 1872; 5 species. LUMBRICLYMENINAE. Nineteen setigers present; nuchal slits strongly curved. Rostrate uncini in single rows; acicular spines present. Anal cone symmetrical, with circular cross-section. Lumbriclymenella Arwidsson 1911a, L. robusta Arwidsson 191 la; only species. LUMBRICLYMENINAE. Nineteen setigers present; nuchal slits strongly curved. Uncini in single rows; acicular spines present. Anal end a simple upturned, flattened cone with anus distal. Macroclymene Verrill 1900, Clymene produca 1897; 2 species. EUCLYMENINAE, More than 30 setigers; slits long and straight. Acicular spines present terior neuropodia. Anal plaque with numerous long cirri. Lewis nuchal in anevenly
Macroclymenella Augener 1926, M. stewartensis Augener 1926; only species, EUCLYMENINAE. More than 30 setigers; nuchal slits long and straight. Acicular spines absent. Setiger 4 with deep encircling collar. Anal plaque with a long midventral and numerous short, slender cirri; anal cone low. Maldane Grube 1860, M. glebifex Grube 1860; 18 species. MALDANINAE. Nineteen setigers; cephalic keel long and high; cephalic rim notched. Acicular spines absent. Maldanella Mclntosh 1885, M. antrctica Mclntosh 1885; 8 species. EUCLYMENINAE. Nineteen setigers; nuchal slits long and straight. Neurosetae absent from first setiger; acicular spines absent. Anal cone large; anal plaque with all cirri evenly short. Microcfymene Arwidsson 1907, M. acirrata Arwidsson 1907; 3 species. EUCLYMENINAE. Nineteen to 20 setigers; nuchal slits long and straight. Acicular spines absent. Anal plaque smooth-rimmed. Micromaldane Mesnil 1897a, M. omithochaeta Mesnil 1897a; 2 species. NICOMACHINAE. Strongly curved, avicular uncini on all setigers; collars absent. Anal plaque with crenulated margin. Nicomache Malmgren 1865, Sabella lumbricalis Fabricius 1780; 16 species.
]976
4 Taxonomic Notes
NICOMACHINAE. Rostrate uncini in single rows; acicular spines in the first three neuropodia. Anal plaque with subequal anal cirri. Notoproctus Arwidsson 1907, N. oculatus Arwidsson 1907; 9 species. LUMBRICLYMENINAE. Nineteen setigers, prostomium large, hoodlike. Acicular spines in anterior neuropodia. Anal plaque flat, smooth-rimmed, with anus dorsal. Petaloprocius Quatrefages 1865, P. terricolus Quatrefages 1865; 11 species. NICOMACHINAE. Nineteen setigers. Acicular spines in the first neuropodia. Anal plaque asymmetrical with the dorsal side strongly reduced. Praxillella Verrill 1881, Praxilla praetermissa Malmgren 1865; 20 species. EUCLYMENINAE. Eighteen to 19 setigers; nuchal slits long and straight. Acicular spines in anterior neuropodia. Anal plaque short, with a long midventral and several shorter anal cirri. Anal cone very large, projecting well beyond the rim of the anal plaque. Praxillura Verrill 1880, P. ornata Verrill 1880; 6 species. LUMBRICLYMENINAE. Twenty to 29 or more setigers; nuchal slits strongly curved. Acicular spines in anterior and posterior neuropodia. Proclymene Arwidsson 1907, Clymene muelleri Sars 1856; only species. EUCLYMENINAE. Nineteen setigers; nuchal slits anteriorly curved. Acicular spines present in anterior neuropodia. Anal plaque with two large midventral and several shorter anal cirri. Pseuoclymene Arwidsson 1907, Clymene quadrilobata Sars 1856; 2 species. EUCLYMENINAE. Eighteen to 20 setigers, nuchal slits short, straight and diverging anteriorly. Acicular spines present; anal cirri all of the same length except the larger midventral one. Rhodine Malragren 1865, R. loveni Malmgten 1865; 8 species. RHODININAE. Numbers of setigers variable; first setigers with notosetae only; acicular spines absent. Numerous encircling collars on posterior setigers. Uncini in double rows. Sonatsa Chamberlin 1919c, S. meridionalis Chamberlin 1919c; only species. MALDANINAE. Numbers of setigers unknown; prostomium with high, long cephalic keel; cephalic rim notched. Rostrate uncini in two or three series in some setigers.
The genus Chaponella Rullier 1972 is considered here in the Sabellidae; it resembles closely members of the genus Euchone, except for its loss of tentacular crown. The genus Sonatsa was described originally in its own subfamily; the posterior end of the only known species remains unknown. The anterior end closely resembles well-known species of Maldane and the genus is considered here in the subfamily Maldaninae. Invalid Genera Arwidssonia Mclntosh 1914b, ses Euclymene Axiothea Malmgren 1865, see. Axiothella Caesicirrus Arwidsson 1911b, set Euclymene Chrysothemis Kinberg 1867b, see Asychis Clymene Oken 1815, indeterminable Clymene Savigny 1818, indeterminable Heteromaldane Ehlers 1908, see Maldane phianissa Kinberg 1867b, questionably PrcwiZ/e/fc Leiocephalus Quatrefages 1865, see Eue lyme ne Leiochone Grube 1868b, see Maldane and Asychis Leiochone Saint-Joseph 1894, see Clymenura Maldanopsis Verrill 1900, &t Asychis Mandrocles Kinberg 1867, indeterminable Neco Kinberg 1867b, see Mylitta Nicomachella Levinsen 1884, see Petaloproctus Paraxiothea Webster 1879a, see Ctymenella Petaloclymene Augener 1918, incompletely known Praxilla Malmgren 1865, see Praxillella Promaldane Mesnil 1897a, hypothetical Sabaco Kinberg 1867b, see Asychis ORDER OPHELIIDA Prostomium without appendages; palps absent. Maximally one anterior asetigerous segment present. Parapodia uniramous or (usually) biramous. All setae simple, including capillary setae. Rostrate hooks absent. All neuropodia short and truncate. Composite setae have been reported from a single species of the family Scalibregmidae; this form may be incorrectly assigned to family and in some respects resembles members of the Syllidae.
FAMILY OPHELIIDAE MALMGREN
1867
Body with a limited number of segments; often with a deep ventral furrow. Prostomium without appendages, blunt or conical. Parapodia biramous or uniramous, with small button-shaped parapodial lobes; all setae capillary, either smooth or marginally dentate. Opheliids have a series of three very distinct bodyshapes. Some are short, thick and grub-shaped; others are very slender, nearly smooth and torpedo-shaped.
42
Science Series 28
Finally, some are anteriorly inflated and posteriorly cylindrical or narrow. Opheliids are common animals
in sandy and muddy bottoms and have been studied extensively by ecologists interested in sandy beaches.
Key to Genera la. 1 b. 2a (la). 2b (la). 3a (2b). 3b (2b). 4a (lb). 4b (lb). 5a (4a). 5b (4a). 6a (4b). 6b (4b). 7a (6a). 7b (6a). 8a (6b). 8b (6b). 9a (8b). 9b (8b). 10a (9b). 10b (9b). Body fusiform without ventral groove 2 Body cylindrical with at least posterior ventrum deeply grooved 4 Branchiae absent Kesun Branchiae present 3 First setiger appear in front of the mouth; all setae smooth Travisia First setiger appear behind the mouth; some setae spinose Dindymenides Ventral groove present in posterior part of body only 5 Ventral groove present along the whole body 6 Three body-regions, including inflated head, inflated anterior part and narrow posterior part; branchiae in posterior region only Euzonus Body not clearly regionated, inflated anteriorly and grooved posteriorly; branchiae from setigers 8-10, if present Ophelia Branchiae absent 7 Branchiae present 8 Lateral eyes absent Tachytrypane Lateral eyes present Polyophtkalmus Lateral eyes present Armandia Lateral eyes absent 9 Anal tube short, all anal cirri of similar length Antiobactrum Anal tube long, two internally attached ventral cirri present, dorsal anal cirri short 10 Branchiae, if present, along the whole body .. Ophelina Branchiae always present, limited to the posterior end of body only Ammotrypanella
Generic Definitions Ammotrypanella Mclntosh 1879, A. rctica Mclntosh 1879; only species. Ventral groove present along the whole body; branchiae present and limited to the posterior end only. Anal tube long and narrow, with two internally attached ventral cirri. Lateral eyes absent. Antiobactrum Chamberlin 1919c, Ophelina brasiliensis Hansen 1882; only species. Ventral groove present along the whole body; branchiae present. Anal tube short and with all anal cirri of the same length. Lateral eyes absent; expanded palpode at the tip of the prostomium. Armandia Filippi 1861, A. cirrhosa Filippi 1861; 18 species. Ventral groove along the whole body; branchiae present. Lateral eyes present; anal tube long and slender, with paired long internally attached ventral cirri and shorter dorsal cirri. Dindymenides Chamberlin 1919c, Dindymene concinna Kinberg 1866b; 3 species. Body short and grnblike without ventral furrow. Branchiae present. First setiger appears behind the mouth; some setae spinose.
FIGURE 12. (A), Family OPHELIIDAE,0;?Mi<i rathkei, modified fix)m Hartmann-Schrder, 1971, about 5x; (B), Family SCALIBREGMIDAE, Scalibregma inflatum, off Anacapa Island, California, 90 m, 5x.
1976
43 Taxonomic Notes
Euzonus Grube 1866, E. arcticus Grube 1866; 10 species. Body with three regions; a head consisting of prostomium and one or two setigers, an anterior swollen region and a posterior region with ventral groove. Branchiae limited to posterior region. Last setiger of anterior region with specialized notopodia. Kesm Chamberlin 1919c, K. fusus Chamberlin 1919c; 5 species. Body short and grublike without ventral furrow. Branchiae absent. Last setigers without epipodial pads; anal cylinder furrowed, but without papillae. Ophelia Savigny 1818, O. bicornis Savigny 1818; 32 species. Body fusiform with inflated anterior end and ventral furrow posteriorly. Branchiae rarely absent, on most setigers from setigers 8-10. No notopodial lobes modified. Ophelina rsted 1843b, O. acuminata rsted 1843b; 44 species. Body with ventral furrow along the whole length. Branchiae presently, rarely absent, on most setigers. Anal cone long, with paired internally attached ventral cirri and shorter dorsal cirri. Lateral eyes absent. Polyophthalmus Quatrefages 1850a, Nais picta Dujardin 1839; 3 species. Body with ventral furrow along the whole length. Branchiae absent; lateral eyes present. Anal tube short with small anal cirri. Tachytrypane Mclntosh 1879, T", jeffreysii Mclmosh 1879; only species. Body with ventral furrow along the whole length. Branchiae and lateral eyes absent. Anal tube cut away ventrally forming an open hood with marginal anal
1 am presently following Hartmann-Schrder (1971) and others in accepting Ophelina as the valid name for worms often referred to under the generic name -4mmotrypane. Hartmann-Schrder (1971) also recognized a series of subfamilies within this family. There are two distinct groups of genera within the family, but the differences between the groups and the size of the family does not seem to warrant such treatment. Invalid Genera Aloysina Clapatde 1864, see Polyophthalmus Ammotrypane Rathke 1843, see Ophelina Cassandane Kinberg 1866b, see Ophelia Dindymene Kinberg 1866b, see Dindymenides Ladice Kinberg 1866b, see Ophelina Nais Dujardin 1839, set Polyophthalmus Neomeris Costa 1844, see Ophelia Nitetis Kinberg 1866b, see Ophelia Omaria Grube 1869b, see Ophelina Pygophelia Sars 1869, indeterminable Terpsichore Kinberg 1866b, see Ophelina Urosiphon Chamberlin 1919c, see Ophelina
FAMILY SCALIBREGMIDAE MALMGREN
1867
Travista Johnston 1840, T. forbesii Johnston 1840, 20 species. Body short and grublike without a ventral groove. Branchiae present; posterior setigers with epipodial pads. Anal cirri all short and thick. All setae smooth; first setiger appears before the mouth.
Body short and stocky or long and slender, often anteriorly inflated. Prostomium anteriorly bifid or with T-shaped frontal horns. Parapodia biramous, with small, triangular or button-shaped parapodia. Setae include capillaries and furcate setae and in some cases acicular spines. Composite setae reported in one species. The scalibregmids contained up to a 'W years ago a ivll-circumscribed group of forms; all characierized by having capillary and furcate setae; usually combined with a strongly wrinkled or areolated epidermis. During last ten years several new forms have been added, making a definition of the family more difficult. Some of these forms {Proscalibregma and Scalispinigera) may in fact not be as closely associated with this family as previously indicated; especially the latter resembles forms usually associated with the order Phyllodocida. No attempts have been made below to move these genera to other families, but it is anticipated that the content of this family may change rather drastically in the near future.
Key to Genera
la. lb. 2a (la). 2b (la). 3a (2b). 3b (2b).
Body short, maggotlike and inflated 2 Body long, sometimes anteriorly inflated, but always with a slender posterior end 4 Two or three anterior asetigerous segments Neolipobranchius One anterior asetigerous segment 3 Branchiae present Polyphysia Branchiae absent Lipobranchius
44 4a (lb). 4b (lb). 5a (4b). 5b (4b). 6a (5b). 6b (5b). 7a (6b). 7b (6b). 8a (7b). 8b (7b). 9a (8a). 9b (8a). 10a (9b). 10b (9b). 1 la (8b). 1 lb (8b). 12a (1 la). 12b (lia). 13a (12b). 13b (12b), 14a (lib). 14b (Ub),
NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY Neurosetae composite heterogomph spinigers Neurosetae capillary, furcate or acicular Furcate setae absent Furcate setae present Median parapodia with long, digitate postsetal lobes Postsetal lobes short and triangular or absent Prostomium with long, cirriform frontal antenna Prostomium T-shaped, without appendages Neither dorsal nor ventral cirri present At least ventral cirri present Acicular setae present Acicular setae absent Eyes present Eyes absent Branchiae present Branchiae absent Acicular spines absent Acicular spines present Eyes present Eyes absent Dorsal cirri absent Dorsal cirri present
Science Series 28 Scalispinigera 5 Proscalibregma 6 Scalibregmides 7 Scalibregmella 8 9 11 Asclerocheilus 10 Hyboscolex Kebuita 12 14 Scalibregma 13 Parasclerocheilus Sclerobregma Sclerocheilus Pseudoscalibregma
Generic Definitions Asclerocheilus Ashworth 1901, Lipobranchius intermedius Saint-Joseph 1894; 4 species. Body elongated, prostomium T-shaped, eyes absent. Qne asetigerous segment. Branchiae, dorsal and ventral cirri absent. Acicular spines in up to three anterior setigers; otherwise with furcate and capillary setae. Hyboscolex Schmarda 1861, H. longiseta Schmarda 1861; 3 species. Body elongated, prostomium T-shaped. Branchiae, dorsal and ventral cirri absent. Setae include capillaries and furcate setae. Eyes present. Kebuita Chamberlin 1919c, Eumenia glabra Ehlers 1887; 2 species. Body elongated, prostomium T-shaped. Branchiae, dorsal and ventral cirri absent. Setae include capillaries and furcate setae. Eyes absent. Lipobranchius Cunningham and Ramage 1888, Eumenia jeffreysii Mclntosh 1869; only species. Body short and maggothke, prostomium bilobed; one asetigerous segment present. Branchiae absent. Acicular setae absent; furcate and capillary setae present. Neolipobranchius Hartman and Pauchald 1971, N. glabrus Hartman and Fauchald 1971; only species. Body short and maggotlike; prostomium a conical lobe. Two or three asetigerous anterior segments present. Acicular spines, furcate setae, branchiae and dorsal and ventral cirri absent. Eyes absent.
Parasclerocheilus Fauvel 1928b, P. branchiatus Fauvel 1928b; 2 species. Body elongated; prostomium T-shaped; eyes present. One asetigerous segment present. Branchiae present. Dorsal cirri absent, ventral cirri present in posterior setigers. Acicular spines present in up to four anterior setigers; other setae include capillaries and furcate setae. Polyphysia Quatrefages 1865, Eumenia crassa rsted 1943b; 2 species. Body short and maggotlike; prostomium bilobed; one asetigerous segment present. Branchiae present. Acicular spines absent; setae include furcate and capillary setae. Proscalibregma Hartman 1967,/*. linea Hartman 1967; only species. Body elongated; prostomium trapezoidal; eyes absent. No asetigerous segment present; all setae capillary; anus within a collar-shaped pygidium, Pseudoscalibregma Ashworth 1901, Scalibregma parva Hansen 1878; 3 species. Body elongated; prostomium T-shaped. One asetigerous segment present. Dorsal and ventral cirri present in posterior setigers. Branchiae absent. Furcate and capillary setae present; acicular spines present in some forms. Scalibregma Rathke 1843, S. inflatum Rathke 1843, 4 species.
976
45 ORDER PHYLLODOCIDA
Body elongated; prostomium T-shaped. Eyes absent. One asetigerous segment present. Branchiae present. Capillary and furcate setae present; acicular spines absent. Scalibregmella Hartman and Fauchald 1971, S. antennata Hartman and Fauchald 1971, only species. Body elongated; prostomium quadrangular with cirriform long antennae and nuchal organs. Dorsal and ventral cirri absent; smooth capillary setae in both rami, furcate setae in notopodia only. A muscular eversible pharynx present. Scalibregmides Hartmann-Schrder 1965, S. chilensis Hartmann-Schrder 1965; only species. Body elongated, prostomium T-shaped, One asetigerous segment present. Branchiae absent. Parapodia with long, slender postsetal lobes in median and posterior setigers. Acicular setae absent, furcate and capillary setae present. Scalispinigera Hartman 1967, S. oculata Hartman 1%7, 2 species. Body elongated; prostomium rectangular; eyes present. One asetigerous segment present. Notosetae all capillary; neurosetae all composite heterogomph spinigers. Branchiae, furcate setae, dorsal and ventral cirri absent. Sclerobregma Hartman 1965, S. branchiata Hartman 1965; only species. Body elongated; prostomium T-shaped, one asetigerous segment present. Eyes present. Branchiae present. Acicular spines in first setiger; capillary and furcate setae present. Sclerocheilus Grube 1863, 5. minutus Grube 1863; 5 species. Body elongated, prostomium T-shaped; one asetigerous segment present. Branchiae absent; ventral cirri present on posterior setigers; dorsal cirri absent. Acicular spines, furcate and capillary setae present. Taxonomic Note The genus Oncoscolex Schmarda 1861, is considered here a synonym of Hyboscolex Schmarda as suggested by Day (1967). Invalid Genera Eumenia Orsted 1843b, see Polyphysia Eumeniopsis Bidenkap 1895, see Polyphysia Eusclerocheilus Hartman 1967, see Pseudoscalibregma Gwasitoa Chamberlin 1919c, see Kebuita Oligobranchus Sars 1846, see Scalibregma Oncoscolex Schmarda 1861, see Hyboscolex
Prostomium usually with at least one pair of antennae; palps, if present, frontal or frontolateral. Maximally two pairs of jaws present, but jaws often absent. Eversible pharynx always muscular and cylindrical, Parapodia distinct in most forms, supported by acicula at least in one ramus. Apart from the amphinomids and eunicids and associated families, this order contains most members of the old order ERRANTIA. It is by far the most speciose of the polychaete orders. For that reason, it was considered appropriate to erect suborders and superfamilies within the order, even if some families are left without specific intermediate designation. Most members of this order are modified moderately from the assumed ancestral condition (Fauchald 1974a) in that they have biramous parapodia and only one, or perhaps a few segments involved in the anterior tagma. Several families show varying levels of cephalization, indicating that this process is going on independently in these taxa. Similarly, a change in function of the notopodia from locomotory to defensive appears to have taken place independently in several of the groups. It is possible to divide this order into several orders recognizing as such, taxa characterized as suborders here. However, the Phyllodocida is characterized by the same characters as all the other orders of polychaetes whereas the suborders of Phyllodocida are characterized by a slightly different set of characters, making such a change undesirable. Suborder PhyUodociformia Phyllodocida with at least two pairs of prostomial appendages and at least two pairs of tentacular cirri. Palps absent; eversible pharynx unarmed; first parapodia lateral.
FAMILY PHYLLOIXJCIDAE WILLIAMS
1851
PhyUodociformia with long and slender bodies; prostomium with four or five antennae, eyes, when present, small. Two to four pairs of tentacular cirri present. Parapodia uniraraous (usually) or biramous with the notopodia represented by a short stalk and large, folise dorsal cirri held erect over the dorsum, rarely with acicula or setae. All neurosetae composite; notosetae, when present, simple. The phyllodocids are common shallow-water polychaetes, more commonly associated with hard substrates than with sands and muds. They are frequently highly colored in life, and these colors are diagnostic, but fade very rapidly upon preservation. The phyllodocids produce copious amounts of mucus when
46
Science Series 28
13. (A), Family ALCIOPIDAE, Vanadis formosa, off central California, pelagic, lOx; (B), seta of the above, 112x; (C), Family PHYLLODOCIDAE, Anaitides groenlandica, Canoe Bay, Alaska, 43-70 m, 12x; (D), seta of the above, lOOx; (E), median parapodium of the above, 19x; (F), Family LOPADORHYNCHIDAE, Lopadorhynchus errans, central Pacific Ocean, deep pelagic, seta from median setiger, 125x; (G), anterior end of the above, iOx; (H), Family PONTODORIDAE, Fontodora pelgica, modified after Uschakov, 1972, about 50x.
FIGURE
976
47
disturbed, and xnust be relaxed carefully before preservation. If possible, each specimen should be preserved separately. Important recent revisions include HartmannSchrder (1971); Uschakov (1972) and Banse (1973). Important identificatory characters and methods of observations: The most important characters in the Phyllodocidae includes the number and arrangement of antennae on the prostomium and the numbers of tentacular cirri and their arrangement on the first few segments (Bergstrm 1914). This latter character may be somewhat troublesome, in that varying fusions obscure the primitive arrangement. In principle, the first segment has a single, dorsal pair of tentacular cirri; the next segment usually both a dorsal and a ventral pair, and the third segment has dorsally a tentacular cirrus and ventrally a normal parapodium. As stated above, discrepancies from this pattern are common, and are of great importance. The best way to observe these structures is to hold the specimen with a pair of forceps so that one can look in at the animal in three-quarter dorsal view, and then rotate the animal slowly from dorsal to lateral positions. In this manner it is possible to follow the segmentai furrows, most easily visible laterally, over towards the dorsal side, and reductions and fusions can be assessed. This whole process is best done under
a stereo microscope, and I have found it helpful to use a low light-angle, so that strong shadows fall across the specimen. In certain groups of species, the ventral tentacular cirri (on the second segment) may be folise, often assymmetrical and flattened. Distortions in fixation of ordinary tapering ventral tentacular cirri may be misleading. The flattening of the ventral cirri is very striking and once seen, this character is never again confused with the distortions one can find in preserved material. The papillation of the eversible pharynx is another important character and if the pharynx is not everted, dissection may be necessary. This is easily done, either ventrally or dorsally on most specimens, but may be difficult to do on smaller animals. However, in most cases it cannot be avoided. Biramous and uniramous parapodia in the phyllodocids are exceedingly similar, and the difference essentially is dened by convention: the parapodia are considered biramous if the notopodia, i.e. the stems of the dorsal cirri, contain internal acicula, or if setae are present. It may be necessary to make parapodial preparations and observe these under the compound microsocpe to ascertain the presence or absence of the internal acicula. The blood-vessels in the notopodial rudiments may resemble the internal acicula, so the observations have to be rather careful.
Key to Genera la. lb. 2a (1 b). 2b (lb). 3a (2a). 3b (2a). 4a (2b). 4b (2b). 5a (4a). 5b (4a). 6a (5b). 6b (5b). 7a (4b). 7b (4b). 8a (7a). 8b (7a). 9a (8b). 9b (8b). lOa (9b). 10b (9b). 11a (7b). 1 lb (7b). 12a (11a). Two antennae present Cirrodoce At least four antennae present 2 Two pairs of tentacular cirri on one segment 3 At least three pairs of tentacular cirri on two or three segments 4 Eversible pharynx diffusely papillated or smooth Eteone Eversible pharynx with large lateral papillae in rows Mysta Three pairs of tentacular cirri on two or three segments 5 Four pairs of tentacular cirri on tiiree segments 7 Third segment with dorsal cirri Lugia Third segment without dorsal cirri 6 Second segment with setae; dorsal and ventral cirri broadly ovate My slides Second segment without setae; dorsal and ventral cirri lanceolate Hesionura Parapodia biramous (with notacicula and sometimes with notosetae) 8 Parapodia uniramous (without notacicula or setae) 11 Ventral tentacular cirrus (on segment 2) foliaceous Hesperophyllum Ventral tentacular cirri digitate or subulate 9 First tentacular segment a complete ring Austrophyllum First tentacular segment dorsally reduced 10 First segment with two pairs of cirri and setae Nipponophyllum First segment with a single pair of cirri; setae first present from the second segment .. Notophyllum Five antenna present 12 Four antennae present 22 Ventral tentacular cirri (on segment 2) folise 13
48 12b (1 la). 13a (12a). 13b ( 12a). 14a (13a). 14b (13a). 15a (13b). 15b (13b). 16a (12b). 16b (12b). 17a (16a). 17b (16a). 18a (16b). 18b (16b). 19a (18b). 19b (18b). 20a (19a). 20b (19a). 21a (I9b). 21b (19b). 22a (lib). 22b (I lb). 23a (22b). 23b (22b). 24a (23a). 24b (23a). 25a (23b). 25b (23b). 26a (25a). 26b (25a). 27a (25b). 27b (25b). 28a (27b). 28b (27b), 29a (28b). 29b (28b). 30a (29b). 30b (29b).
Science Series 28
Ventral tentacular cirri digitate or subulate 16 First segment dorsally reduced 14 All tentacular segments complete rings 15 Second tentacular segment with setae Sige Second tentacular segment without setae Pterocirrus Second tentacular segment with setae; proboscis papillated Steggoa Second tentacular segment without setae; proboscis smooth Natalia Nuchal epaulettes present 17 Nuchal epaulettes absent 18 First and second tentacular segment free from each other and complete rings Vitiazia First and second tentacular segment fused; first segment dorsally reduced Vitiaziphyllum First and second tentacular segment fused; pharynx smooth Eumida All three tentacular segments free from one another; pharynx papillated 19 Reniforra ventral cirri broadly attached transverse to the long axis of the parapodia 20 Ventral cirri narrowly attached; usually narrow and elongated 21 Pharyngeal papillae distally in six rows Bergstroemia Pharyngeal papillae dispersed Clavadoce Pharynx with few papillae; second segment with setae Pirakia Pharynx with numerous papillae; second segment without setae Eulalia All three tentacular segments form complete rings Protomystides and Pseudeulalia (these two cannot be separated on current information). First two segments fused dorsally 23 Nuchal papillae absent 24 Nuchal papillae present 25 Tentacular cirri flattened Nereiphylla Tentacular cirri digitate or subulate Genetyllis First and second segment fused to each other and to the prostomiura 26 First and second segment fused to each other at least dorsally, but free from the prostomium .... 27 Simple enlarged setae in segments 2-4 Chaetoparia Segments 2-4 with composite spinigers Prochaetoparia First segment with a pair of small dorsolateral papillae in addition to the tentacular cirri Prophyllodoce First segment with tentacular cirri only 28 First to third pair of tentacular cirri and the antenna globose, four pairs of cirri digitate Sphaerodoce All pairs of tentacular cirri and the antennae digitate or subulate 29 Nuchal papilla on a posterior prolongation from the prostomium Paranaitis Prostomiura heart-shaped with the nuchal papilla in a posterior incision 30 Pharyngeal papillae basally in rows Anaitides Pharyngeal papillae basally dispersed Phyllodoce
Taxonomic Notes The system indicated above, follows in the main the principles first promulgated by Bergstrm (1914) and followed by most later authors. It incorporates the main features of the systems organized by Uschakov (1972) and Banse (1973). Certain taxa here considered genera often are considered as subgenera. This creates a problem of setting the relative taxonomic value of the same character in different parts of the family, to the subgeneric level in some cases and to the generic level in others (cf. Banse 1973). For the sake of consistency, these characters are here all considered to be of generic value.
Generic Dfinitions Anaitides Czemiavsky 1882, Phyllodoce groenlandica rsted 1843a; 28 species. Phyllodocids with four antennae; heart-shaped prostomium with nuchal papilla in the crevice between the lobes. First segment dorsally reduced and partially fused to the second; all tentacular cirri cylindrical, Papillae in rows on the base of the eversible pharynx, Parapodia uniramous. Austrophyllum Bergstrm 1914, Eulalia charcoti Gravier 191 lb; 5 species, Phyllodocids with five antennae; pentagonal or
1976
49
rounded prostomium and without a nuchal papilla. All anterior segments complete and free from one another; all tentacular cirri cylindrical. Parapodia blramous. Bergstroemia Banse 1973, Eulalia nigrimaculata Moore 1909; only species. Phyllodocids with five antennae, pentagonal prostomium and without a nuchal papilla. All anterior segments free from one another, but the first dorsally reduced; all tentacular cirri cylindrical. Eversible pharynx with papillae in six rows distally. Parapodia uniramous, ventral cirri large and reniform and attached transverse to the long axis of the parapodium. Chaetoparia Malmgren 1867, C. nilssoni Malmgren 1867; only species, Phyllodocids with four antennae and a nuchal papilla; prostomium fused to the first segment, which, in turn, is fused to the second. All tentacular cirri cylindrical. Specialized, enlarged simple setae in segments 2-4. Cirrodoce Hartman and Fauchald 1971, C. cristate Hartman and Fauchald 1971; only species. Phyllodocids with two antennae, quadrangular prostomium and without nuchal organs. Three pairs of tentacular cirri on two segments; first segment with normal setae. A pair of auricular lobes attached between prostomium and first parapodia. Clavadoce Hartman 1936c, C. splendida Hartman 1936c; 2 species. Phyllodocids with five antennae, quadrangular prostomium and no nuchal papilla. First segment dorsally reduced, but free from the other tentacular segments; all four pairs of tentacular cirri club-shaped and slightly flattened. Eversible pharynx with diffuse papillation. Parapodia uniramous, ventral cirri large and reniform and attached transverse to the long axis of the parapodium. Eteone Savigny 1818, Nereis flava Fabricius 1780; 40 species. Phyllodocids with four antennae; triangular or trapezoidal prostomium and a small nuchal papilla. All tentacular segments complete and free from one another; two pairs of cylindrical tentacular cirri present. Eversible pharynx diffusely papillated or smooth. Parapodia uniramous. Eulalia Savigny 1818, Nereis viridis Linnaeus 1767; 50 species. Phyllodocids with five antennae; pentagonal prostomium and no distinct nuchal papilla. All three tentacular segments free from one another and from the prostomium; all tentacular cirri cylindrical. Eversible pharynx diffusely papillated; parapwdia uniramous. Eumida Malmgren 1865, Eulalia sangunea rsted 1843b; 13 species.
Phyllodocids with five antennae; pentagonal prostomium and indistinct nuchal papilla. First tentacular segment dorsally reduced and fused to the second; all tentacular cinri cylindrical. Eversible pharynx smooth or with very small papillae. Parapodia uniramous. Genetyllis Malmgren 1865, G. lutea Malmgren 1865; 11 species. Phyllodocids with four antennae; short wide prostomium and no nuchal papilla. First and second tentacular segments are fused and reduced dorsally; all tentacular cirri cylindrical. Eversible pharynx diffusely papillated. Parapodia uniramous. Hesionura Hartmann-Schrder 1858, H. fragilis Hartmann-Schrder 1958; 9 species. Phyllodocids with four antennae, prolonged prostomium and no nuchal papilla. Two tentacular segments, free from each other and from the prostomium; all three pairs of tentacular cirri cylindrical. Third segment without dorsal cirrus. Hesperophyllum Chamberiin 1919a, /i. tectum Chamberlin 1919a; only species. Phyllodocids with five antennae, short and wide prostomium and no nuchal papilla. All tentacular segments complete; ventral cirrus foliaceous; all others cylindrical. Parapodia blramous; ventral cirrus large and transversely attached. Lugia Quatrefages 1865, Eteone aurantiaca Schmarda 1861; 3 species. Phyllodocids with five antennae of which the median is very small; long prostomium and no nuchal papilla. Two tentacular segments with three pairs of tentacular cirri; all cylindrical. Third segment with dorsal cirrus. Parapodia uniramous. Mysta Malmgren 1865, M. barbota Malmgren 1865; 6 species. Phyllodocids with four antennae; pentagonal prostomium and no nuchal papilla. Two pairs of tentacular cirri on first segment; all cirri cylindrical. Eversible pharynx with large lateral papillae in rows. Mystides Thel 1879, M. borealis Thel 1879; 8 species. Phyllodocids with five antennae; long prostomium and no nuchal papillae. Three pairs of cylindrical tentacular cirri on two segments. Third segment without dorsal cirri. Nereiphylla Blainville 1828, N. paretti Blainville 1828; 3 species. Phyllodocids with four antennae; short wide prostomium and no nuchal papilla. First and second segment fused and dorsally reduced; all tentacular cirri flattened. Eversible pharynx diffusely papillated. Parapodia uniramous; ventral cirrus reniform.
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Nipponophyllum Imajima and Hartman 1964, Notophy Hum japonic urn Marenzeller 1879; 3 species. Phyllodocids with five antennae; rounded pentagonal prostomium and no nuchal papilla. Four pairs of cylindrical tentacular cirri on two segments; first segment dorsally reduced, but with setae. Eversible pharynx closely papillated. Parapodia biramous. Notalia Bergstrm 1914, Eulalia picta Kinberg 1866b, only species. Phyllodocids with five antennae; pentagonal prostomium and no nuchal papilla. All tentacular segments free from one another and from the prostomium; ventral cirri folise; all others cylindrical. Eversible pharynx smooth; parapodia uniramous, setae present from second segment. Notophyllum rsted 1843b, Pkyllodoce foliosa Sars 1835; 6 species. Phyllodocids with five antennae; broadly transverse prostomium and nuchal epaulettes present. All tentacular segments free, but first or first and second dorsally reduced; all tentacular cirri cylindrical. Parapodia biramous; no setae in first segment. Paranaitis Southern 1914, Anaitis wahlbergi Malmgren 1865; 15 species. Phyllodocids with four antennae; rounded prostomium and a nuchal papilla present. First and second segment fused and dorsally reduced; all tentacular cirri cylindrical. Eversible pharynx with rows of lateral papillae. Parapodia uniramous. Pkyllodoce Savigny 1818, P. laminosa Savigny 1818; 48 species. Phyllodocids with four antennae; heart-shaped prostomium with nuchal papilla in the crevice between the lobes. First segment dorsally reduced and fused to the second; all tentacular cirri cylindrical. Eversible pharynx with diffuse papillation. Parapodia uniramous. Pirakia Bergstrm 1914, Pkyllodoce {Eulalia) punctifera Grube 1860; 4 species. Phyllodocids with five antennae; pentagonal prostomium and no nuchal papilla. All three anterior segments fi-ee from each other; all tentacular cirri cylindrical. Eversible pharynx with few scattered papillae. Parapodia uniramous; setae present from second segment. Prochaetoparia Bergstrm 1914, Genetyllis brevis Ehlers 1901; only species. Phyllodocids with four antennae; broadly transverse prostomium; a nuchal papilla present. First and second segment fused to each other and to the prostomium; all tentacular cirri cylindrical. Parapodia uniramous, with normal composite setae in all setigers. Prophyllodoce Hartman 1966b, P. hawaiia Hartman 1966b; only species.
Phyllodocids with four antennae; heart-shaped prostomium with a nuchal papillae between the lobes. First and second segment fused; first segment dorsally reduced; all tentacular cirri cylindrical. First segment with a pair of small papillae dorsal to the tentacular cirri. Prostomium with diffuse papillae. Parapodia uniramous. Protomystides Czemiavsky 1882, Mystides bidentata Langerhans 1880; only species. Phyllodocids with four antennae and no nuchal papilla. All tentacular segments complete rings; three or four pairs of cylindrical tentacular cirri on three segments. Pseudeulalia Eliason 1962, P. exigua Eliason 1962; only species. Phyllodocids with four antennae and no nuchal papilla. All tentacular segments are separate and complete rings; all tentacular cirri are cylindrical. Parapodia uniramous. Pterocirrus Claparde 1868, Pkyllodoce (Eulalia) macToceros Grube 1860; 10 species. Phyllodocids with five antennae; pentagonal prostomium and no nuchal papilla. All tentacular segments free from one another; first segment dorsally reduced; ventral tentacular cirrus folise, all others cylindrical. Parapodia uniramous; setae first present in second segment. Sige Malragren, 1865, Sige fusigera Malmgren, 1865; about five species. Phyllodocids with five antennae, penagonal prostomium and no nuchal papilla. All tentacular segments free from one another; first segment dorsally reduced; ventral tentacular cirrus folise, all others cylindrical Parapodia uniramous, setae first present in the third segment. Sphaerodoce Bergstrm 1914, Phillodoce quadraticeps Grube 1878, only species. Phyllodocids with four antennae; heart-shaped prostomium with a nuchal papilla between the lobes. First and second segments fused and dorsally reduced; first three pairs of tentacular cirri and the antennae globose; last pair of tentacular cirri (on third segment) digitate. Steggoa Bergstrm 1914, Eulalia magalaensis Kinberg 1866b; 13 species. Phyllodocids with five antennae, pentagonal prostomium and no nuchal papilla. All three tentacular segments complete rings, fiêe from one another. Ventral tentacular cirrus folise; all others cylindrical, Eversible pharynx diffusely papillated. Parapodia uniramous. Vitiazia Uschakov 1953, V. dogieli Uschakov 1953; only species.
1976
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Phyllodocids with four large and one small median antenna; broadly truncate prostomium and paired nuchal epaulettes. All tentacular segments fully developed and separate from one another; tentacular cirri cylindrical. Parapodia uniramous. Vitiaziphyllum Uschakov 1972, V. nuchalum Uschakov 1972; only species. Phyllodocids with five antennae, broadly transverse prostoinium and paired nuchal epaulettes. First and second segment fused; first segment reduced dorsally; all tentacular cirri cylindrical. Parapodia uniramous.
Invalid Genera Carobia Quatrefages 1865, see Nereiphylla Carobia (Paracarobia) Czemiavsky 1882, see Anahides Carobia (Protocarobia) Czemiavsky 1882, see Phyllodoce Cirraria Sveshnikov 1959, larval forms Eracia Quatrefages 1865, see Eulalia ana Sige Eteonella Mclntosh 1874, see Eteone Eteonides Hartmann-Schrder 1960a, see Hesionwa Eulalides Czemiavsky 1882, see Eumida (?) Eumenia Quatrefages 1865, error for Eunomia, indeterminable Eumidia Verriil 1873b, error for Eumida Eunomia Risso 1826, indeterminable Eunotophyllum Czemiavsky 1882, sec Notophyllum Globiodoce Bergstrm 1914, lapsus calami ioxSphaerodoce Hypereteone Bergstrm 1914, see Eteone Hypocirrus Giard 1913, indeterminable
Hypoeulalia Bergstrra 1914, see Eulalia Kinbergia Quatrefages 1865, indeterminable Lepadorhynchus Schmarda 1861, see Anaitides Macrophyllum Schmarda 1861, see Notopkyllum Mesoeulalia Czemiavsky 1882, indeterminable Mesomystides Czemiavsky 1882, see My slides Myriana Savigny 1820, indeterminable Myriacycium Grube 1880, see Eulalia Nothis Pruvot 1885, hypothetical Paraeulalia Czemiavsky 1882, see Notophyllum Phyllodoce Ranzani 1817, see Polyodontes (family Polyodontidae) Porroa Quatrefages 1865, see Eulalia Protocarobia Czemiavsky 1882, see Phyllodoce Pseudonotophyllum Czemiavsky 1882, see Notophyllum Trachelophyllum Levinsen 1883, see Notophyllum
FAMILY ALCIOPIDAE EHLERS
1864
Phyllodociforms with slender, transparent long bodies found exclusively pelagically. Prostomium with five antennae and a pair of very large, spherical eyes. Three to five pairs of tentacular cirri. Parapodia uniramous, dorsal and ventral cirri foliaceous; setae simple or composite. Alciopids are delicate, slender pelagic organisms, mainly known for their very large, complex cameratype eyes (Hermans and Eakin 1974). The pelagic polychaetes recently were reviewed by Dales and Peter (1972). The system indicated below, follows these two authors. The key to genera has been rewritten from Uschakov (1972).
Key to Genera lalb. 2a (la). 2b (la). 3a (2b). 3b (2b). 4a (lb). 4b (lb). 5a (4b). 5b (4b). 6a (5a). 6b (5a). 7 (5b). 7b (5b). 8a (7b). 8b (7b). Several anterior segments with mdimentary parapodia 2 Anterior segments with fully developed parapodia 4 All setae simple capillaries Naiades Setae composite spinigers 3 Parapodia distally with a cirriform appendage Vanadis Parapodia distally without a cirriform appendage Torrea Parapodia with two distal cirriform appendages, nearly all setae composite spinigers Alciopa Parapodia without cirriform appendages, or with a single such appendage; simple or acicular setae present 5 All setae simple g At least some composite setae present 7 Parapodia with a cirriform appendage Krohnia Parapodia without cirriform appendages Alciopina Parapodia with cirriform appendages 8 Parapodia without cirriform appendages Plotohdmis Acicula barely extending beyond the tip of the parapodia Rhynchonerella Aciculum prolonged, extending well beyond the tip of the parapodia Watelio
52
NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY Generic Definitions
Science Series 28
Alciopa Audouin and Milne Edwards 1833b, A. reynaudi Audouin and Milne Edwards 1833b; one, possibly three species. Alciopids with parapodia fully developed on the anterior segments. Each parapodium with two digitiform cirri distally. Three or four pairs of tentacular cirri. Setae include numerous composite spinigers, but a few slender capillaries may be present. Alciopina Claparde and Panceri 1867, A. parasitica Claparde and Panceri 1867; 2 species. Alciopids with parapodia fully developed on anterior segments. Parapodia without digitiform cirri, Five pairs of tentacular cirri. All setae simple, including numerous slender capillaries and several shorter acicular setae, Krohnia Quatrefages 1865, Alciopa lepidota Krohn 1845; 2 species. Alciopids with parapodia fully developed on anterior segments. Parapodia with a single digitiform cirrus distally. Five pairs of tentacular cirri. All setae simple, including capillaries and thicker, acicular setae in anterior segments and ventrally in other parapodia. Naiades delle Chiaje 1830, N. cantrainii delle Chiaje 1830; only species. Alciopids with rudimentary parapodia on first three body segments. Three pairs of tentacular cirri. Parapodia without digitiform cirrus. All setae simple capillaries. Plotohelmis Chamberiin 1919c, Rhynchonerella capitata Greeff 1876; 3 species. Alciopids with parapodia fully developed on anterior segments. Four or five pairs of tentacular cirri present, Parapodia without digitiform distal cirrus. Setae of two kinds including simple acicular setae and composite setae with slender pointed appendages. Rhynchonerella Costa 1862, R. gracilis Costa 1862; 5 species. Alciopids with parapodia fully developed on anterior segments. Five pairs of tentacular cirri present. Parapodia with single digitiform distal cirri. Setae of several kinds, including slender composite setae with long appendages, and shorter acicular setae simple or composite. Torrea Quatrefages 1^5, Alciopa candida delle Chiaje 1828; 2 species.
Alciopids with rudimentary parapodia on the two first segments. Three pairs of tentacular cirri. Parapodia without digitiform distal cirri. All setae composite spinigers with long narrow appendages. Vanadis Claparde 1870b, V. formosa Claparde 1870b; 13 species. Alciopids with several segments with rudimentary parapodia, maximally ten segments. Three or four pairs of tentacular cirri. Each parapodium with a digitiform distal cirrus. All setae composite spinigers with long, slender appendages. Watelio Stp-BowiU 1948, Callizona gravieri Benham 1929; only species. Alciopids with parapodia fully developed on all segments. Four pairs of tentacular cirri present. Each parapodium with a digitiform distal cirrus; acicula extending well beyond the tip of the parapodia. Setae include numerous composite spinigers with long, slender appendages and a few simple capillaries. Invalid Genera Asterope Claparde 1870b, ste. Torrea Callizona Greeff 1876, stt Rhynchonerella CaUizonella Apstein 1891, see. Krohnia Cleta Claparde 1870b, lapsus calami for Vanadis Corynocephalus Levinsen 1885, ste. Alciopina Greeffia Mclntosh 1885, ste Alciopa Halodora Greeff 1876, see Alciopa Liocapa Costa 1862, see Torrea Mauita Chamberiin 1919c, see Vanadis Nauphanta Greeff 1876, see Alciopa 1868
FAMILY LOPADORHYNCHIDAE CLAPARDE
Exclusively pelagic, short-bodied phyllodociform polychaetes. Prostomium with four antennae and a pair of small eyes. Two or three pairs of tentacular cirri present. Parapodia uniramous, dorsal and ventral cirri cirriform; all setae composite except for modified anterior setae in some forms. Members of this family often are considered as truly phyllodocids and are treated as such. It is preferable to segregate them in their own family; inclusion in the phyllodocids would make that family very difficult to define. Dales and Peter (1972) and Uschakov (1972) have given recent reviews; but the latter considered the lopadorhynchids as part of the phyllodocids.
Key to Genera la. lb. 2a (la). Setae present at the base of the tentacular cirri Setae absent at the base of the tentacular cirri Two pairs of tentacular cirri on one segment; parapodial cirri long and digitate 2 3 Pelagobia
1976 2b (la). 3a (lb). 3b (lb).
53 Maupasia Lopadorhynchus Pedinosoma
Three pairs of tentacular cirri on two segments; parapodial cirri short First two or three parapodia modified and with large, simple setae All parapodia similar, all setae composite spinigers
Generic Definitions Lopadorhynchus Grube 1855, L, brevis Grube 1855, 9 species. Lopadorhynchids with three pairs of tentacular cirri on two segments; setae absent from tentacular segments. Two or sometimes three anterior setigers enlarged with markedly modied parapodia and simple setae. Normal segments with conical setal lobes and rounded presetal lobes. Normal setae composite with broad, oar-shaped appendages with dentate cutting margin. Maupasia Viguier 1886, M. caeca Viguier 1886, 4 species. Lopadorhynchids with three pairs of tentacular cirri on two segments; setae present on tentacular segments. Al! parapodia similar; each conical with short dorsal and ventral cirri. Setae include composite spinigers with slender appendages and sometimes simple setae in the first seliger. Pedinosoma Reibisch 1895, P. curium Reibisch 1895, only species. Lx)padorhynchids with two pairs of tentacular cirri on the first segment; no setae on tentacular segment. All parapodia similar, with small rounded dorsal cirri and long, slender ventral cirri. All setae composite spinigers. Pelagobia Greeff 1879, P. longicirrata Greeff 1879, 2 species. Lopadorhynchids with two pairs of tentacular cirri on first segment; setae present on tentacular segment. All parapodia similar with long setal lobes and very long dorsal and ventral cirri. All setae composite with narrowly oar-shaped, marginally dentate appendage. Invalid Genera Haiiplanella Treadwell 1943, ite Maupasia Haiiplanes Reibisch 1895, see. Maupasia Halyplanes Reibisch 1893, see Maupasia Hydrophones Claparde 1870b, sts Lopadorhynchus
Mastigethus Chamberlin 1919c, &s Lopadorhynchus Reibischia Bergstrm 1914, xt Lopadorhynchus

FAMILY PONTODORIDAE BERGSTRM
1914
Phyllodociform, small, slender, pelagic polychaetes. Prostomium with four antennae, eyes small. Two pairs of tentacular cirri present. Slender, elongated uniramous parapodia with long dorsal cirri; setae composite. The family is known for a single genus, Pontodora Greeff 1879 and a single species P. pelgica Greeff 1879. It has so far been reported from warm waters in the Atlantic Ocean and from colder waters in the Pacific Ocean (Uschakov 1972). Invalid Genus Epitoka Treadwell 1943, see Pontodora Suborder Aphroditiformia Phyllodocida with up to three antennae (may be absent); one pair of tentacular cirri; long ventral palps. Eversible pharynx, if armed, with two or four jaws. First parapodia directed forwards. Superfamily Aphroditacea Dorsal ciiri alternating with elytrae in at least part of the body; notosetae cylindrical in cross-section, slender or thick. Prostomium not fused to the first setiger.
FAMILY APHRODITIDAE MALMGREN
1867
Aphroditaceans with dorsoventrally flattened bodies and one antenna present. Eversible pharynx with a pair of jaws or unarmed. Elytrae alternating with dorsal cirri; each marginally smooth or irregularly lobed. Notosetae of several kinds, including in some genera finely attenuated felt setae and in others harpoonshaped, erect, flattened spines or cylindrical spines. All setae simple.
Key to Genera la. lb. 2a (la). 2b (la). 3a (2b). 3b (2b). Dorsum covered with felted notosetae Dorsum without feh, or with a very few feh setae Stout supportive notosetae absent; facial tubercle absent Stout supportive notosetae present; facial tubercle present Neutosetae distally bifid Neurosetae distally entire, but sometimes subdistally spurred 2 4 Heteraphrodita 3 Tricertia Aphrodita
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Science Seres 28
FIGURE
14. Family APHR0DrnDAE,/4pr>t//ia refulgida, Halfmoon Bay, California, felt removed on right side, 2.5x.
4a (lb). 4b (lb). 5a (4b). 5b (4b). 6a (5b). 6b (5b).
Harpoon-shaped notosetae present Harpoon-shaped notosetae absent Notosetae sabre-shaped and smooth Notosetae, if flattened, marginally serrated Notosetae flattened and marginally serrated Notosetae cylindrical and smooth
Laetmonice 5 Aphrogenia 6 Pontogenia Hermionopsis
Generic Definitions Aphrodita Linnaeus 1758,^4. aculeata Linnaeus 1761; 26 species. Facial tubercle present; felt present. Harpoon-shaped notosetae absent; other long supportive notosetae present. Aphrogenia Kinberg 1855, A. alba Kinberg 1855; 5 species. Facial tubercle present; felt absent. Harpoon-shaped notosetae absent; notosetae long, sabrelike or smooth and capillary. Hermionopsis Seidler 1923, H. levisetosa Seidler 1923; only species. Facial tubercle present; felt absent. Harpoon-shaped notosetae absent; notosetae long, straight and cylindrical without asperities. Heteraphrodita Pettibone 1966a, H. altoni Pettibone 1966a; only species. Facial tubercle absent; felt present. Stout supportive notosetae absent. Laetmonice Kinberg \i55, L. filicomis Kinberg 1855; 28 species. Facial tubercle present; felt absent. Harpoon-shaped notosetae present. Pontogenia Claparde 1868, Hermione chrysocoma Baird 1865; only species. Facial tubercle present; felt poorly developed or absent. Protective notosetae flattened and marginally serrated. Tricertia Haswell 1883, T. araeoceras Haswell 1883; only species. Facial tubercle present; felt present. Long protective notosetae present, flattened and curved. Neurosetae distally bifid.
1976 Taxonomic Notes
55
The above treatment follows Pettibone (1966a) closely in that several genera (e.g. Hermonia Hartman 1959), generally considered valid, have been excluded from the named genera. The validity oiTricertia and Hermionopsis is doubtful; however, both have been defined on characters usually used in the group, and therefore, have been included. Invalid Genera Bartulara Sveshnikov 1959^ NOMENNUDUM Cyanippa Castelnau 1842, s&eAphrodita
Halithea Savigny 1818, s&t Laetmonice Halogenia Horst 1916b, sec Laetmonice Hermione Blainville 1828, stc Laetmonice Hermonia Hartman 1959, st^ Laetmonice Laetmatonice Ex A UCTORE, see Laetmonice Letmonicella Roule 1898, ste Laetmonice Miinesia Quatrefages 1865, ste Aphrodita
FAMILY POLYNOIDAE MALMGREN
1867
Aphroditaceans with dorsoventrally flattened bodies; one, two or, usually, three antennae present. Eversible pharynx with four jaws. Marginally smooth or fringed
FIGURE
IS. (A), Family POLYNOIDAE, Halosydna brevisetosa, Sunset Bay, Oregon, intertidal, elytrae removed on left side, tOx; (B), Family POLYODONTIDAE, Polyodomes sp-, off Santa Barbara, California, 84 m, elytrae removed on the right side, 15x.
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elytrae alternate with dorsal cirri at least on the anterior end. All setae simple, notosetae variously developed, but never as felt- or harpoon-setae, and usually distinctly lateral in position. The polynoids are the most commonly occurring scale-worms. They are usually medium-sized, rarely large worms. Forms with smooth elytrae are often commensals with other organisms; those with strongly ornamented elytrae tend to be free-living. Nearly 100 genera of scale-worms presently are recognized and more are being described every year; most genera are well defined and the species are recognizable, but within several of the larger genera (Harmothoe, Lepidonotus, Halosydna) problems occur. Several taxa that usually are considered as subgenera under Harmothoe, have been considered as separate genera
below. They differ from Harmothoe SENSV STRICTU, in exactly the characters that usually are considered valid generic characters elsewhere in the family and cannot be related to Harmothoe by overlap. If possible, when preserving scale-worms, especially members of HARMOTHOINAE, care must be taken that scales are maintained with the specimens, since these furnish one of the best diagnostic characters for species identifications. The best way of doing this, is either to bulk-relax the whole sample (MS 222, or MgClj) or, if the specimens are larger, preserve each scale-wotm separately. There are no recent reviews of the whole family; recent authors interested in these worms, include Hartman and Hartmann-Schrder, and especially Marian H. Pettibone of the Smithsonian Institution.
Key to Genera 1 a. lb. 2a (lb). 2b (lb). 3a (2a). 3b (2a). 4a (3a). 4b (3a). 5a 5b 6a 6b (2b). (2b). (5b). (5b). Cervilia 2 3 MACELLICEPHALINAE 5 Median antenna present 4 Median antenna absent IPHIONINAE 7 Lateral antennae attached sub-distally or ventrally on the prostomium (ceratophores below the anterior points of the prostomium which may be produced into cephalic peaks) HARMOTHOINAE 8 Lateral antennae attached distally on the prostomium (ceratophores continuations of the prostomium) LEPIDONOTINAE 51 Fifteen pairs of elytrae Macelloides Maximally 13 pairs of elytrae 6 Prostomium consisting of two inflated lobes, 16-17 segments present Macellicephaloides Prostomium a single unified lobe, with paired rounded anterior projections, 17-19 segments Macellicephala Notosetae coarser than neurosetae Bylgides Notosetae much thinner than neurosetae phione First setigers with simple, strongly curved hooks 9 Anterior hooks absent 10 Two segments with anterior hooks Australaugeneria Three segments with anterior hooks Uncopolynoe Fourteen or fewer paks of elytrae 11 Fifteen or more pairs of elytrae 15 Notosetae absent Polynoella Notosetae present 12 Eight or nine pairs of elytrae present, parapodia prolonged Herdmanella Twelve to 14 pairs of elytrae present, parapodia short 13 Some notocirri with spherical subdistal inflations Andresia Notocirri maximally gently inflated 14 Eyes very large and confluent Robertianella Eyes small Antinoana Fifteen pairs of elytrae 16 Sixteen or more pairs of elytrae 42 More than 50 segments present, only anterior part of body covered by elytrae 17 Less than 45 segments present; elytrae cover the whole body, or at most, the last 10-15 segments uncovered 23 Lateral antennae inserted posteriorly on the prostomium Lateral antennae inserted anteriorly At least two antennae well developed Median antenna well developed; lateral antennae reduced or absent
7a (3b). 7b (3b). 8a (4a). 8b (4a). 9a (8a). 9b (8a). lOa (8b). 10b (8b). 1 la (10a). 1 lb ( 10a). 12a ( I lb). 12b (lib). 13a (12b). 13b (12b). 14a (13b). 14b (13b). 15a (10b). 15b (10b). 16a (15a). 16b (15a).
]<)76
17a ( 16a). 17b 16a). 18a ( 17b). 18b 17b). 19a ( 18b). 19b 18b). 20a { 19a). 20b 19a). 21a 19b). 21b 19b). 22a 21b). 22b 21b).
51
23a 23b 24a 24b 25a 25b 26a 26b 27a 27b 28a 28b 29a 29b 30a 30b 31a 31b 32a 32b 33a 33b 34a 34b 35a 35b 36a 36b 37a 37b 38a 38b 39a 39b 40a 40b 41a 41b 42a 42b 43a 43b 44a 44b
16b). 16b). 23a). 23a). 24b). 24b). 25b). [25b). 23b). (23b). 27b). (27b). 28b). (28b). 29b). (29b). 30b). (30b). 31a). (31a). 31b). (31b). 33a). (33a). 34b). (34b). 35b). (35b). 33b). (33b). 37b). (37b). 38b). (38b). 39b). (39b). 40b). (40b). 15b). (15b). (42a). (42a). 43b). (43b).
Neurosetae with semilunar pockets Adyte Neurosetae spinose or smooth, without pockets 18 Notosetae absent Heteropolynoe Notosetae present 19 All notosetae thicker than or at least as thick as the neurosetae 20 At least some capillary notosetae present 21 Notosetae held erect over body Hermadion Notosetae short and lateral in position Hemilepidia Eyes reduced or absent, prostomium quadrangular Nemidia Eyes distinct, prostomium rounded 22 Notosetae mainly capillary, neurosetae unidentate Enipo Notosetae mainly thick and blunt-tipped, neurosetae bidentate, except in far posterior setigers Polynoe Notosetae distinctly slenderer than the neurosetae 24 At least some notosetae as thick as or thicker than the neurosetae 27 Ventral surface with paired folise appendages Phyllohartmania Ventral surface smooth 25 Most neurosetae bidentate Arcteobia All neurosetae unidentate 26 Neurosetae slender, straight spines Hartmania Neurosetae distally falcate Gattyana Ventral cirri folise, ventrum papillose Phyllosheila Ventral cirri digitate, ventrum usually smooth 28 Neurosetae in part distally trifid Gorekia Neurosetae distally uni- or bidentate 29 Neurosetae with semilunar pockets Paradyte Neurosetae spinose or nearly smooth 30 Notosetae with distal tufts of long hairs Barrukia Notosetae without distal tufts of hairs 31 Neurosetae distally with capillary tips 32 Neurosetae distally falcate, uni- or bi-dentate 33 Some neurosetae with fine, filamentous tips Antinoella All neurosetae with slender tips, but without filaments Antinoe All neurosetae unidentate 34 At least some neurosetae bidentate 37 Dorsal cirri alternating between short and long; short dorsal cirri basally inflated .... Kermadecella All dorsal cirri similar in length, none basally inflated 35 Notosetae of two kinds, thick and slender Hesperonoe All notosetae thick and similar in shape 36 Eyes absent Intoshella Eyes present , Eunoe Neurosetae distally pencillate Austrolaenilla Neurosetae never pencillate 38 Both noto- and neurosetae slender; notosetae with capillary tips Tenonia Notosetae and most neurosetae coarse 39 Neurosetae include furcate spines with subequal tips and slender, dentate capillaries Melaenis Neuropodial capillary setae absent 40 Some neurosetae deeply cleft with both teeth long and narrow Eucranta Neurosetae with main fangs very much larger than secondary teeth 41 Posterior eight to ten segments not covered by elytrae Lagisca Whole body covered by elytrae Harmothoe Sixteen pairs of elytrae 43 Eighteen or more pairs of elytrae 45 Neurosetae with semilunar pockets Subadyte Neurosetae slender and serrate, semilunar pockets absent 44 Notosetae distinctly serrated Leucia Notosetae with a few spines Scalisetosus
58 45a (42b). 45b (42b). 46a (45b). 46b (45b). 47a (46a). 47b (46a). 48a (47b). 48b (47b). 49a (46b). 49b (46b). 50a (49b). 50b (49b). 51a (4b). 51b (4b). 52a (51b). 52b 53a 53b 54a 54b 55a 55b 56a (51b). (52b). (52b). (53b). (53b). (54b). (54b). (55b).
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56b (55b). 57a (56b). 57b (56b). 58a (57a). 58b (57a). 59a (58a). 59b (58a). 60a (59b). 60b (59b). 61a (58b). 61b (58b). 62a (61b). 62b (61b). 63a (62b). 63b (62b). 64a (63b). 64b (63b). 65a (64b). 65b (64b). 66a (57b). 66b (57b). 67a (66a). 67b (66a). 68a (66b), 68b (66b). 69a (68b). 69b (68b). 70a (69a). 70b (69a). 71a (70b). 71b (70b).
Eighteen pairs of elytrae Acanthicolepis Nineteen or more pairs of elytrae 46 Notosetae distinctly slenderer than neurosetae 47 Notosetae as thick as, or thicker than neurosetae 49 Neurosetae with semilunar pockets Pottsiscalisetosus Semilunar pockets absent 48 Neuropodia with subacicular digitiform lobes Parahotalepidella Neuropodia with large, supra-acicular presetal lobes Grubeopolynoe All neurosetae unidentate Polyeunoa At least some neurosetae bidentate 50 Neurosetae with long bare tips beyond the serrated regions Hololepidella Neurosetae with very short bare tips beyond the serrated region Neohololepidella Palps represented by two small tubercles Bouchiria Palps well developed 52 Ventral cirrus heavily fimbriated, ventrum with transverse rows of three papillae on each segment Lepidofimbria Ventral cirrus and ventrum smooth 53 Neurosetae distally bifid with several small accessory teeth in the crotch Lepidogyra Neurosetae unidentate or bifid, rarely trifid; if bifid, then without accessory teeth 54 Prostomium anteriorly produced into two large lobes with small lateral antennae attached distally Allmaniella Prostomium anteriorly tapering into the ceratophores 55 Renal papillae greatly prolonged in some median setigers Bathymoorea Renal papillae of similar length in all setigers 56 Notopodia completely reduced, neuropodium prolonged with the ventral cirrus attached near the middle Frennia Notopodia usually present; if absent, then neuropodia short with ventral cirri basal 57 Twelve or 13 pairs of elytrae 58 More than 13 pairs of elytrae 66 With branchial filaments on the elytrophores 59 Branchial filaments absent 61 With pseudelytrae on non-elytrophoral, alternating segments posteriorly Euphionella Pseudelytrae absent 60 Neurosetae coarsely spinose Chaetacanthus Neurosetae finely plumose Euphione Notopodia absent Drieschia Notopodia present 62 With one very large bidentate hook in the middle of each neuropodial fascicle Sheila All neurosetae similar in thickness, none bidentate hooks 63 Neurosetae distally trifrcate Hermenia Neurosetae distally entire 64 Notosetae of two kinds, lancet-shaped and tapering and serrated Thormora Notosetae all of one kind, usually slender and serrated 65 Pseudelytrae on non-elytrophoral segments Dilepidonotus Pseudelytrae absent Lepidonotus Fourteen pairs of elytrae 67 Fifteen or more pairs of elytrae 68 Neurosetae bidentate Lucopia Neurosetae entire Podarmus Only anterior half of body covered by elytrae Pseudopolynoe Elytrae on most parts of the body 69 Fifteen pairs of elytrae 70 Sixteen or more pairs of elytrae 74 Notosetae thin Parahalosydna Notosetae as thick as, or thicker than the neurosetae 71 All neurosetae unidentate Eulagisca At least some neurosetae bidentate 72
]976 72a (71b). 72b (71b). 73a (72b). 73b (72b). 74a (69b). 74b (69b). 75a (74b). 75b (74b). 76a (75a). 76b (75a). 77a (76a). llh (76a). 78a (77b). 78b (77b). 79a (76b). 79b (76b). 80a (75b). 80b (75b). 81a (80a). 81b (80a). 82a (81b). 82b (81b). 83a (82b). 83b (82b). 84a (83a). 84b (83a). 85a (84b). 85b (84b). 86a (83b). 86b (83b). 87a (86a). 87b (86a). 88a (86b). 88b (86b). 89a (88b). 89b (88b). 90a (80b). 90b (80b). 91a (90b). 91b (90b). 92a (91a). 92b (91a). 93a (91b). 93b (91b).
59
Dorsal cirri of two kinds, short and expanded or long and gently inflated Malmgreniella All dorsal cirri long and slightly inflated subdistally 73 Notosetae nearly smooth Malmgrenia Notosetae densely serrated Paralepidonotus With 16 pairs of elytrae Arctonoella At least 17 pairs of elytrae 75 Seventeen to 20 pairs of elytrae 76 At least 21 pairs of etytrae 80 Notosetae absent 77 Notosetae present 79 Dorsum with a median pustule on each segment Weheria Dorsal pustules absent 78 Neurosetae with capillary tips Drieschella Neurosetae with falcate tips Alentiana Notosetae numerous, lateral antennae terminal Haiosydna Only few notosetae present; lateral antennae subterminal Alentia Notosetae present 81 Notosetae absent 90 Parapodia distally with series of long papillae Halosydnopsis Parapodia distally without papillae 82 Neurosetae with semilunar pockets Hololepida Neurosetae spinose or smooth 83 Notosetae thinner than most neurosetae 84 Notosetae as thick as, or thicker than the neurosetae 86 Superior neurosetae slender and spinose, inferior ones acicular Pseudohalosynda All neurosetae stout 85 At least some neurosetae bidentate Halosydnella All neurosetae unidentate Acholoe All neurosetae unidentate 87 At least some neurosetae bidentate 88 Ventrum with large segmentally arranged lamellae Gastrolepidia Ventrum smooth Lepidastheniella Lateral antennae subterminal, neurosetae few Arctonoe Lateral anteimae terminal, neurosetae numerous 89 Twenty-one or 22 pairs of elytrae; neurosetae with long, curved tips Hyperhalosydna Numerous pairs of elytrae; neurosetae with short, straight or curved tips Lepidasthenia Ventral cirri with two-three knobs Perolepis Ventral cirri smooth 91 Antennal scales present 92 Antennal scales absent 93 Setae flattened, marginally serrated with long, slender tips; up to 30 pairs of elytrae Admetella Setae flattened, finely dentate and very abruptly tapering; 23 pairs of elytrae Bathyadmetella Neurosetae stout, falcate; pre- and postsetal neuropodial lobes of similar length . .. Benhamipolynoe Neurosetae slender; presetal lobes distinctly longer than the postsetal ones in the neuropodia Telolepidasthenia Generic Definitions LEPIDONOTINAE. Numerous segments and elytrae. Cirriferous segments with T-shaped papillae. Notosetae thinner than neurosetae, with transverse rows of spines. Neurosetae few, unidentate with serrated subdistal areas. Admetella Mclntosh 1885, A. longipedata Mclntosh 1885; 2 species. LEPIDONOTINAE. Up to 30 pairs of elytrae; 75 segments. Antennal scales at base of lateral antennae. Notosetae absent; neurosetae long, flattened, finely
Acanthicolepis Mclntosh 1900, Polynoe asperrima Sars 1861; 3 species. HARMOTHOINAE. Eighteen pairs of elytrae, approx. 40 segments. Notosetae thicker than neurosetae, with whorls of teeth. Neurosetae bidentate with slender secondary tooth; serrated. Dorsal cirri and antennae papillated. Acholoe Claparde 1870b, Nereis squamosa delle Chiaje 1828; only species.
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marginally serrated with smooth tips. Neuropodia prolonged. Adyte Saint-Joseph 1899, Hertnadion assimile Mclntosh 1876; only species. HARMOTHOINAE, Fifteen pairs of elytrae; long posterior region without scales. Notosetae at least as thick as neurosetae, smooth with few spines. Neurosetae with semilunar pockets and faintly bifid tips; serrations faint. Presetal neuropodial lobes longer than postsetal ones. Alenda Malmgren 1865, Polynoe gelatinosa Sars 1835; 3 species. LEPIDONOTINAE. Eighteen pairs of elytrae; 43 segments. Lateral antennae subterminal. Notopodia reduced with a few, nearly capillary setae. Neurosetae thin, slender, usually bifid. Alentiana Hartman 1942, Polynoe aurantiaca Verrill 1885; only species, LEPIDONOTINAE. Seventeen to 20 pairs of elytrae; 36-39 segments. Notosetae absent. Neurosetae unidentate, serrated or smooth. Allmaniella Mclntosh 1885, A. setubalensis Mclntosh 1885; 5 species. LEPIDONOTINAE. More than 15, but less than 30 pairs of elytrae. Prostomium anteriorly produced into two large lobes with the small lateral antennae attached distally. Notosetae thick and finely serrated; neurosetae of two kinds, upper slender and smooth, lower thicker, bidentate and vaguely serrated. Andresia Prenant 1924, A. ampullifera Prenant 1924: only species. HARMOTHOINAE. Thirteen to 14 pairs of elytrae 32-33 segments. Notosetae thicker than neurosetae coarsely serrated. Neurosetae unidentate, serrated Some notocirri with large, spherical ampullae subdistally. Antinoana Hartman and Fauchald 1971, A. fusca Hartman and Fauchald 1971; only species. HARMOTHOWAE. Twelve to 13 pairs of elytrae; 26-27 segments. Notosetae thicker than neurosetae, vaguely serrated. Neurosetae slender and bifid. Antinoe Kinberg 1855, A. microps Kinberg 1855; 8 species. HARMOTHOINAE. Fifteen pairs of elytrae. Notosetae thicker than neurosetae, blunt, transversely serrated. Neurosetae with slender tips, but not Glamentously prolonged. Antinoella Augener 1928b, Antinoe sarsi Kinberg in Malmgren 1865; 7 species. HARMOTHOINAE. Fifteen pairs of elytrae; approximately 40 segments. Notosetae thicker than neuro-
setae with rows of spines. Neurosetae distally with slender, filamentous tips, often pilose. Arcteobia Annenkova 1937, Eupolynoe anticostiensis Mclntosh 1874; 2 species. HARMOTHOINAE. Fifteen pairs of elytrae; 36 segments. Notosetae slenderer than neurosetae, with blunt or capillary tips. Upper neurosetae sharp-tipped; others bidentate. Arctonoe Chamberlin 1920, Polynoe vittata Grube 1855; 4 species, LEPIDONOTINAE. Forty or more pairs of elytrae, numerous segments. Lateral antennae subterminal. Notosetae few, thick, recurved uni- or bidentate with coarse serrations. Neurosetae few, thick and falcate, uni- or bidentate. Arctonoella Buzhinskaya 1967, Harmothoe sinagawaensis Izuka 1912; only species. LEPIDONOTINAE. Sixteen pairs of elytrae, 41 segments. Lateral antennae subterminal. Notosetae thinner than neurosetae, distally capillary. Neurosetae unidentate. Australaugeneria Pettibone 1969d, Polynoe rutilans Grube 1878; 2 species. HARMOTHOINAE, Fifteen pairs of elytrae, approximately 40 segments, Notosetae thicker than neurosetae, smooth or faintly serrated. Neurosetae varied: as large golden hooks in the second and third segment; otherwise uni- or bidentate, often spurred and subdistally inflated. Austrolaenilla Bergstrm 1916, A. antrctica Bergstrm 1916; 6 species. HARMOTHOINAE. Fifteen to 16 pairs of elytrae; 40-43 segments. Notosetae thicker than neurosetae with transverse rows of teeth, Neurosetae unidentate or bidentate, with the distal end pencillate. Barrukia Bergstrm 1916, Gattyana cristata Willey 1902; 2 species. HARMOTHOINAE. Fifteen pairs of elytrae, 35-36 segments. Notosetae distally with tuft of long, fine hairs, otherwise blunt and serrated, Neurosetae unidentate, with a few coarse teeth on the cutting edge. Bathyadmetella Pettibone 1967, B. commando Pettibone 1967; only species. LEPIDONOTINAE. Twenty-three pairs of elytrae, 58 segments. Notosetae absent, notopodia reduced. Neuropodia prolonged, with slender, flattened setae, finely dentate and distally very abruptly tapering. Bathymoorea Pettibone 1967, Polynoe Trenotuberculata Moore 1910; only species. LEPIDONOTINAE, Numbers of elytrae not known, 35 segments. Notosetae short and slender, neurosetae
1976
61
thicker, long, bidentate and marginally serrated. Renal papillae greatly prolonged in some median setigers. Benhamipolynoe Pettibone 1970d, Lepidasthenia antipathicola Benham 1927; only species. LEPIDONOTINAE. More than 17 pairs of elytrae; numerous segments. Notosetae absent. Neurosetae thick, falcate with weakly marked serrations. Bouchiria Wesenberg-Lund 1949, B. vesiculosa Wesenberg-Lund 1949; only species. LEPIDONOTINAE. Nimibers of pairs of elytrae and segments not known. Notosetae absent; neurosetae include slender capillary setae with long slender spines and thicker more acicular, but otherwise similar setae ventrally. Palps reduced to two small tubercles. Numerous stalked papillae present on the parapodia. Bylgides Chamberlin 1919c, Bylgia elegans Thel 1879; only species. IPHIONINAE. Two antennae; facial tubercle absent. Notosetae coarser than neurosetae. Cervilia Frickinger 1916, C. japnica Frickinger 1916; only species. Sub-family unknown. Three antennae, lateral antennae attached posteriorly on the prostomium. Fifteen pairs of elytrae. Notosetae slender and capillary; neurosetae unidentate and thicker. Chaetacanthus Seidler 1924, Iphione magnifica Grube 1875; 3 species. LEPIDONOTINAE. Twelve pairs of elytrae, 26 segments. Elytrophores with branchiae. Notosetae fine and capillary; neurosetae unidentate and spinose. Dilepidonotus Hartman 1967, D. falklandicus Hartman 1967; only species. LEPIDONOTINAE. Twelve pairs of elytrae, 26 setigers. Notosetae slender and silky capillaries; neurosetae thicker, smooth and pointed. Dorsum of cirral segments crested; pseudelytrae present. Drieschia Michaelsen 1892, D. pelgica Michaelsen 1892; 6 species. LEPIDONOTINAE. Twelve to 13 paire of elytrae; 26-27 segments. Notopodia absent; upper neurosetae capillary, lower thicker and slightly serrated. Drieschella Augener and Pettibone in Pettibone 1970d, D. maculata Augener and Pettibone in Pettibone 1970d; only species. LEPIDONOTDJAE. Twenty pairs of small elytrae; 47 segments. Notosetae absent; neurosetae slender and tapering to capillary tips. Presetal lobes longer than postsetal ones; acicular lobes not projecting. Enipo Malmgien 1865, E. kinbergi Malmgren I86S; 4 species.
HARMOTHOINAE. Fifteen pairs of elytrae; posterior half of body not covered by elytrae. Notosetae mainly capillaries, a few thicker blunt setae present. Neurosetae thicker than notosetae, mainly unidentate, but a few bidentates also present. Prostomium round, eyes conspicuous. Eucranta Malmgren 1865, E. villosa Malmgren 1865; 4 species. HARMOTHOINAE. Fifteen pairs of elytrae; 36^0 segments. Notosetae thicker than neurosetae with rows of teeth. Some neurosetae distally split with both parts about equally long and thick; other neurosetae slender and unidentate. Eulagisca Mclntosh 1885,. corrientis Mclntosh 1885; 3 species. LEPIDONOTINAE. Fifteen pairs of elytrae; 37 segments. Lateral antennae subterminal; facial tubercle present, Notosetae slender, numerous, pectinate, thicker than the neurosetae. Neurosetae slender and tapering to capillary tips. Eunoe Malmgren 1865, E. oerstedi Malmgren 1865; 40 species. HARMOTHOINAE. Fifteen pairs of elytrae, approximately 40 segments. Notosetae thicker than neurosetae with rows of spines; neurosetae all unidentate, with more or less well-marked spinose region. Euphione Mclntosh 1885, E. elisabethae Mclntosh 1885; 6 species. LEPIDONOTINAE. Twelve pairs of elytrae, shortbodied. Elytrophores with branchiae. Notosetae capillary; neurosetae thicker, unidentate and laterally covered with fine hairs. Euphionella Monro 1936, Physalidonotus lobulatus Seidler 1922; 3 species, LEPIDONOTINAE. Twelve pairs of elytrae; 25 segments. Elytrophores with branchiae. Notosetae capillary; neurosetae thicker, unidentate and smooth. Pseudelytrae present, Frennia Viguier 1912, F. dubia Viguier 1912; 2 species. LEPIDONOTINAE. Numbers of pairs of elytrae and segments not known. Notopodia completely reduced; neuropodia strongly prolonged with ventral cirri attached near the middle. All setae slender, smooth capillaries. Gastrolepidia Schmarda 1861, G. clavigera Schmarda 1861; only species. LEPIDONOTINAE. More than 21 pairs of elytrae. Notosetae and neurosetae similar in thickness; notosetae blunt and serrated; neurosetae unidentate and serrated. Anteiuiae and dorsal cirri strongly inflated subdistally, with slender tips, Ventrum with large lamellae.
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Gattyana Mclntosh 1900, Aphrodita cirrhosa Pallas 1766; 14 species. HARMOTHOINAE. Fifteen pairs of elytrae; approximately 40 segments. Most notosetae capillary with fine dentition. Neurosetae thicker than notosetae, distally unidentate, serrated. Gorekia Bergstrm 1916, Malmgrenia crassicirris Willey 1902; only species. HARMOTHOINAE. Fifteen pairs of elytrae; 38-40 segments. Notosetae shorter and thicker than the neurosetae, finely serrated. Neurosetae in part distally trifid. Grubeopolynoe Pettibone 1969b, Polynoe tuta Grube 1855; only species. HARMOTHOINAE. Fifty or more pairs of elytrae; numerous segments. Notosetae slenderer than neurosetae; of two kinds, short and blunt and slender and tapering, Neurosetae all of one kind, vaguely bidentate or unidentate, serrated. Neuropodia with long supraacicular postsetal lobe. Halosydna Kinberg 1855,//. patagnica Kinberg 1855; 14 species. LEPIDONOTINAE. Eighteen pairs of elytrae; 37 segments. Notosetae much fmer than neurosetae, pointed and serrated. Neurosetae thick, uni- or bidentate, with coarse serrations. Halosydnella Hartman 1938, Halosydna australis Kinberg 1855; 9 species, LEPIDONOTINAE. Twenty-one pairs of elytrae; 45 segments. Notosetae fmer than neurosetae; serrated. Neurosetae distally uni- or bidentate, subdistally serrated. Halosydnopsis Uschakov and Wu 1959, Halosydna pilosa Horst 1917; only species. LEPIDONOTINAE. Twenty-seven pairs of elytrae, body covered. Notosetae fmer than neurosetae, finely serrated; neurosetae nearly smooth and unidentate. Distal end of parapodia with series of large papillae. Harmothoe Kinberg 1855, H. spinosa Kinberg 1855; 120 species. HARMOTHOINAE. Fifteen pairs of elytrae; approximately 40 segments. Notosetae thicker than neurosetae; with rows of spines. Neurosetae at least in part bidentate, but usually also some unidentate in inferior positions. Hartmania Pettibone 1955, H. moorei Pettibone 1955; only species. HARMOTHOESTAE. Fifteen pairs of elytrae; 40 segments. Notosetae ner than neurosetae, tapering to sharp tips. Neurosetae tapering to sharp, pointed tips, not falcate. Hemilepidia Schmarda 1861, . tuberculata Schmarda 1861; 4 species.
HARMOTHOINAE. Fifteen pairs of elytrae; posterior part of body not covered by elytrae. Notosetae as thick as neurosetae, vaguely serrated. Neurosetae distally bi- or unidentate. Herdmanella Darboux 1899, Polynoe ascidioides Mclntosh 1885; 3 species. HARMOTHOINAE. Eight to 9 pairs of elytrae; 15-17 segments. Notosetae spinose, neurosetae less so, both conspicuously long and unidentate, parapodia prolonged. Hermadion Kinberg 1855, H. magalhaensi Kinberg 1855; 5 species. HARMOTHOINAE. Fifteen pairs of elytrae; more than 50 segments. Notosetae thicker than neurosetae and strongly serrated. Neurosetae uni- or bidentate, serrated. Notosetae held erect over body. Hermenia Grube 1856, H. verruculosa Grube 1856; 2 species. LEPIDONOTINAE. Twelve pairs of elytrae, small, not overlapping. Notosetae few, slender and serrated; neurosetae distally trifrcate. Hesperonoe Chamberlin 1919, Harmothoe complnala Johnson 1901; 4 species. HARMOTHOINAE, Fifteen pairs of elytrae; 36-38 segments. Notosetae in part at least as thick as neurosetae; of two kinds; thick serrated and blunt and slender, serrated and pointed. Neurosetae all unidentate, and serrated, usually slender superior and thick in inferior positions. Heteropolynoe Bidenkap 1907, H. nordgaardi Bidenkap 1907; only species. HARMOTHOINAE, Numbers of pairs of elytrae not known, 58 segments, Notosetae absent, all neurosetae unidentate and marginally serrated, slender in superior and coarse in inferior positions, Hololepida Moore 1905, H. magna Moore 1905; 3 species. LEPIDONOTINAE. Numerous pairs of elytrae, numerous segments, Notosetae few in numbers, capillary; neurosetae thicker, of two kinds, lanceolate and bidentate with transverse rows of pockets. Nuchal flap present. Hololepidella Willey 1905, H. commensalis Willey 1905; 5 species. HARMOTHOINAE. Twenty-six or more pairs of elytrae; 55 or mote segments. Notosetae at least as thick as neurosetae, nearly smooth with blunt tips. Neurosetae distally bidentate or entire. Hyperhalosydna Augener 1922, Lepidonotus striatus Kinberg 1855; 2 species. LEPIDONOTINAE. Twenty-one to 22 pairs of elytrae; 50 segments. Notosetae few, short, curved
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and blunt. Neurosetae bidentate with long, curved tips. Lateral antennae terminal. Intoshella Darboux 1899, Polynoe (Langerhansia) euplectellae Mclntosh 1885; 3 species. HARMOTHOINAE. Fifteen pairs of elytrae, approximately 40 segments. Noto- and neurosetae similar in thickness; notosetae smooth-tipped, weakly serrated; neurosetae unidentate, more distinctly serrated. Eyes absent. Iphione Kinberg 1855, Polynoe muricata Savigny 1818, 4 species. IPHIONINAE. Thirteen pairs of elytrae; notosetae capillary, neurosetae serrated and distally entire. Kermadecella Darboux 1899, Polynoe magnipalpa Mclntosh 1885; only species. HARMOTHOINAE. Fifteen pairs of elytrae, short body. Notosetae thicker than neurosetae, serrated. Neurosetae with transverse rows of spines; distally unidentate. Dorsal cirri alternating long and short, the short ones basally inflated. Lagisca Malmgren 1865, Polynoe rarispina Sars 1861; 24 species. HARMOTHOINAE. Fifteen pairs of elytrae, approximately 50 segments. Noto- and neurosetae about equally thick. Notosetae with dense rows of teeth; neurosetae at least in part bidentate; all neurosetae serrated. Posterior 8-10 segments not covered by elytrae. Lepidasthenia Malmgren 1867, Polynoe elegans Grube 1840; 37 species. LEPIDONOTDslAE. Numerous pairs of elytrae and segments. Notosetae few, usually blunt; neurosetae numerous, uni- or bidentate, most are thicker than notosetae except inferior ones in each fascicle. Lateral antennae terminal. Lepidastheniella Monro 1924, Polynoe comma Thomson 1902; 3 species. LEPIDONOTESIAE. Up to 90 pairs of elytrae; covering the body. Notosetae thinner than neurosetae, ringed with spines. Neurosetae spinose and distally entire. Lepidofimbria Hartman 1967, L. oculata Hartman 1967; only species. LEPIDONOTINAE. Numbers of pairs of elytrae and segments not known. Ventrum with transverse row of three papillae on each segment; ventral cirrus heavily fimlniated. Notosetae aosent; neurosetae smooth and distally entire. Lepidogyra Hartman 1967, L. alba Hartman 1967; only species. LEPIDONOTINAE. Numbers of pairs of elytrae and segments not known. Notosetae coarser than neurosetae; neurosetae spinose, distally bind with a series
of accessory teeth in the crotch between the two major teeth. Lepidonotus Leach 1816, Aphrodita clava Montagu 1808; 65 species. LEPIDONOTINAE. Twelve pairs of elytrae; 26 segments. Notosetae finer than neurosetae, all tapering with whorls of spines. Neurosetae with rows of coarse teeth; rarely bidentate, usually unidentate. Leucia Malmgren 1867, Polynoe nivea Sars 1863; only species. HARMOTHOINAE. Sixteen pairs of elytrae, shortbodied. Notosetae coarser than neurosetae and serrated. Neurosetae long, slender, unidentate and serrated. Lucopia Pillai 1965, L. magnicirra Pillai 1965; only species. LEPIDONOTINAE. Fourteen pairs of elytrae; 27 segments. Notosetae absent; neurosetae bidentate and serrated. Dorsal cirri strongly inflated. Macellicephala Mclntosh 1885, M. mirabilis Mclntosh 1885; 18 species. MACELLICEPHALINAE. Eight to 13 pairs of elytrae; 17-29 segments. Notosetae few or absent, if present then with marginal teeth. Neurosetae long, usually paddle-shaped. Macellicephaloides Uschakov 1955 b, M. grandicirra Uschakov 1955b; only species. MACELLICEPHALINAE. Maximally 9 pairs of elytrae; 16-17 segments. Notosetae absent. Neurosetae long, marginally dentate. Prostomium with two very strongly inflated lobes. Macelloides Uschakov 1957, M. antrctica Uschakov 1957; only species, MACELLICEPHALINAE, Fifteen pairs of elytrae; 30 segments. Notosetae absent. Neurosetae distally inflated. Malmgrenia Mclntosh 1874, M. whiteavesi Mclntosh 1874; 9 species. LEPIDONOTINAE. Fifteen pairs of elytrae; 36-41 segments. Noto- and neurosetae similar in thickness. Notosetae nearly smooth; neurosetae uni- or bidentate with very small secondary teeth. Lateral antennae subterminal. Maimgreniella Hartman 1967, M. dicirra Hartman 1967; only species. LEPIDONOTINAE. Fifteen pairs of elytrae; 41-56 segments. Noto- and neurosetae similar in thickness; notosetae falcate, nearly smooth. Neurosetae bidentate with long, slender secondary tooth. Dorsal cirri of two kinds; long and slender and short and expanded. Melaenis Malmgren 1865, M. lovent Malmgren 1865; only species.
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HARMOTHOINAE. Fifteen pairs of elytrae; 39-41 segments. Notosetae thicker than neurosetae, few in number; smooth or faintly structured. Neurosetae of two kinds: numerous slender dentate with capillary tips; few furcates with subequal blunt tips. Nemidia Malmgren 1865, N. torelli Malmgren 1865; 9 species. HARMOTHOINAE. Fifteen pairs of elytrae; posterior part of body without elytrae. Notosetae mainly capillaries, but a few thick spines present; neurosetae mainly unidentate, but a few bidentate setae present. Prostomium quadrangular, eyes missing or strongly reduced. Neohololepidella Pettibone 1969b, A', murrayi Pettibone 1969b; only species. HARMOTHOINAE. Fifty or more pairs of elytrae; numerous segments. Notosetae thicker than neurosetae, nearly smooth, with blunt tips. Neurosetae with very short bare tip beyond a dense spinose region; distally bidentate or entire. Paradyte Pettibone 1969a, Polynoe crinoidicola Potts 1910; 2 species. HARMOTHOINAE. Fifteen pairs of elytrae; 40 segments. Notosetae thicker than neurosetae, sabrelike with entire or slightly notched tips, nearly smooth. Neurosetae of two kinds: supracicular ones with semilunar pockets, slender, spinose and with bifid tips. Subacicular ones thicker, with semilunar pockets and entire tips. Parahalosydna Horst 1915a, P. sibogae Horst 1915a; 4 species. LEPIDONOTINAE. Fifteen pairs of elytrae; shortbodied. Notosetae thinner than neurosetae, serrated. Neurosetae unidentate and serrated along both edges. Parahololepidella Pettibone 1969b, Hololepidella greeffii Augener 1918; 2 species. HARMOTHOINAE. Numerous pairs of elytrae and segments. Notosetae slenderer than neurosetae, but still thick. Neurosetae very thick, slightly hooked, entire and very faintly spinose. Neuropodia with subacicular digitiform process. Paralepidonotus Horst 1915a, Polynoe ampullifera Grube 1878; 4 species. LEPIDONOTDJAE. Fifteen pairs of elytrae; 38 segments. Notosetae thicker than the neurosetae; densely serrated. Neurosetae at least in part bidentate. Perolepis Ehlers 1908, P. regularis Ehlers 1908; only species. LEPIEXDNOTINAE. Numerous pairs of elytrae and segments. Notosetae absent; neurosetae distally bidentate. Ventral cirri with three knobs; cirrophores
enlarged dorsally; pre- and postsetal lobes of similar length. Phyllohartmania Pettibone 1961, P. taylori Pettibone 1961; only species. HARMOTHOINAE. Fourteen pairs of elytrae, less than 40 segments. Notosetae slender and spinose with capillary tips; neurosetae similar in thickness; distally spinose and spinigerous. Ventral surface with paired folise appendages on each segment. Phyllosheila Pettibone 1961, P. wg/fyi Pettibone 1961; only species. HARMOTHOINAE. Fifteen pairs of elytrae; less than 50 segments. Notosetae thicker than neurosetae, spinose. Neurosetae smooth and distally bidentate. Ventral cirri folise, ventral surface papiUated. Podarmus Chamberlin 1919c, P. pica Chamberlin 1919c; 2 species. LEPIDONOTINAE. Fourteen pairs of elytrae, 30 segments. Notosetae absent; neurosetae all distally entire; of two kinds, thick and straight or slender and capillary. Polyeunoa Mclntosh 1885, P. laevis Mclntosh 1885; 3 species. HARMOTHOINAE. Nineteen to 30 pairs of elytrae, posterior part of body not covered by elytrae. Notosetae thicker than neurosetae, faintly serrated. Neurosetae unidentate, subdistally expanded and dentate. Polynoe Savigny 1818, P. scolopendrina Savigny 1818; 17 species. HARMOTHOINAE. Fifteen pairs of elytrae; posterior part of body not covered by elytrae. Notosetae mainly thick and blunt-tipped, but a few capillary setae present. Nearly all neimisetae bidentate, except usually one or two unidentate in each of the posterior setigers; most neurosetae coarser than the notosetae. Polynoella Mclntosh 1885, P. levisetosa Mclntosh 1885; 3 species. HARMOTHOINAE. Twelve pairs of elytrae; 25-26 segments. Notosetae absent. Neurosetae long, slender, unidentate and falcate. Neuropodia long, pointed and distally bifid. Pottsiscalisetosus Pettibone 1969a, Scatisetosus praelongus Marenzeller 1902; only species. HARMOTHOINAE. Twenty-eight or more pairs of elytrae; nmnerous segments. Notosetae finer than neurosetae; tapered to blunt tips, serrated. Neurosetae with semilunar pockets, distally entire and more or less falcate. Pseudohalosydna Fauvel 1913, P. rosea Fauvel 1913; only species. LEPIDONOTINAE. At least 20 pairs of elytrae; numerous segments. Notosetae spinose and capillary;
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neurosetae of two kinds; superior ones slender and spiralled spinose; inferior ones acicular and spinose. Pseudopolynoe Day 1962, Polynoe inhacae Day 1951; only species. LEPIDONOTINAE. Fifteen to 17 pairs of elytrae; posterior half of body not covered by elytrae. Notosetae finer than neurosetae, serrated; neurosetae bi- or unidentate, serrated. Robertianella Mclntosh 1885, R. synophthalma Mclntosh 1885; only species. HARMOTHOINAE. Thirteen pairs of elytrae; approximately 30 segments. Noto- and neurosetae of similar thickness; notosetae blunt, nearly smooth; neurosetae distally minutely notched, nearly smooth. Eyes very large and nearly confluent on the prostomium. Scalisetosus Mclntosh 1885, S. ceramensis Mclntosh 1885; 8 species. HARMOTHOINAE. Sixteen pairs of elytrae; 40 segments. Notosetae much coarser than neurosetae, with a few spines only. Neurosetae slender, distally entire and spinose. Both noto- and neuropodia with long lobes. Sheila Monro 1930, S. bathypelagica Monro 1930; only species. LEPIDONOTINAE. Thirteen pairs of elytrae; 29 segments. Notosetae slender, dentate capillaries; neurosetae of several kinds: numerous superior dentate capillaries; most of the others coarse, dentate and entire; one very large bidentate hook in the middle of each fascicle. Subadyte Pettibone 1969a, Polynoe pellucida Ehlers 1864; 3 species. HARMOTHOINAE. Fifteen to 16 pairs of elytrae; approximately 40 segments. Notosetae similar in thickness to the neurosetae; with spinose pockets and slightly notched tips. Neurosetae with semilunar pockets, spinose and distally bidentate. Telolepidasthenia Augener and Pettibone in Pettibone 1970d, T. lobetobiensis Augener and Pettibone in Pettibone 1970d; only species. LEPIDONOTINAE. More than 21 pairs of elytrae and more than 50 segments. Notosetae absent; neurosetae slender with spinose regions and entire tips. Presetal lobes longer than postsetal ones. Tenonia Nichols 1969, T. kitsapensis Nichols 1969; only species. HARMOTHOINAE. Fifteen pairs of elytrae; approximately 40 segments. All setae similar in thickness; all slender; notosetae capillary, neurosetae in part bidentate. Thormora Baird 1865, T.jukesii Baird 1865; 5 species. LEPIDONOTINAE. Twelve pairs of elytrae; short-
bodied. Notosetae of two kinds; short and lancetshaped, and longer, serrated and tapering. Neurosetae serrated and unidentate. Uncopolynoe Hartmann-Schrder 1960a, /. comllicola Hartmann-Schrder 1960a; only species. HARMOTHOINAE. Numbers of elytrae not known, approximately 44 segments. Notopodia absent; first three neuropodia with strongly curved hooks; others with uni- or bidentate setae with serrated subdistal areas. Weberia Horst 1915b, W. pustulata Horst 1915b; 3 species. LEPIDONOTINAE. Eighteen pairs of elytrae; shortbodied. Notosetae absent; neurosetae unidentate and curved. Ventral cirri absent except in second segment; dorsum with a pustule on each segment.
Invalid Genera Agnodice Mclntosh 1885, see Lagisca Bathynoe Ditlevsen 1917, see Weberia Bylgia Thel 1879, see Bylgides Chaetosphaera Haecker 1898, larvae of several genera Dasylepis Malmgren 1867, see Acanthicolepis Eumolpe Oken 1807, see Lepidonotus Eupolynoe Mclntosh 1874, see Eucrunw Evarne, Malmgren 1865, see Harmothoe Evarnella Chamberlin 1919c, see Harmothoe Gastroceralella Darboux 1899, êe Thormora Halosydnoides Seidler 1924, see Arctonoe Harmopsides Chamberlin 1919c, see Lepidasthenia Iphionella Mclntosh 1885, seelphione Laenilla Malmgren 1865, see Harmothoe Langerhansia Mclntosh 1885, see Intoshella Lepidametria Webster 1879b, see Lepidasthenia Nectochaeta Marenzeller 1892, in part Lepidasthenia, also generally polynoid larvae and juveniles Norepa Baird 1865, seelphione Nychia Malmgren 1865, see Gattyana Oligolepis Levinsen 1887, see Macellicephala Paranychia Czemiavsky 1882, questionably laîico Parapolynoe Czemiavsky 1882, see Polynoe Parmensis Malmgren 1867, see Harmothoe Pkysalidonotus Ehlers 1905, see Euphione Plotolepis Chamberlin 1919c, see Drieschia Quetieria Viguier 1911, juvenile form Tricosmochaeta Morgera 1918, see Harmothoe
FAMILY POLYODONTIDAE BUCHANAN
1894
Aphroditaceans with dorsoventrally flattened bodies. Two or three antennae present, median antenna when
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present, attached dorsally or posteriorly on the prostomium (antennae may be absent). Eversible pharynx with four jaws. Marginally fringed or smooth elytrae alternate with dorsal cirri in all setigers. All setae simple. Spinning glands present.
The polyodontids are large, solid-bodied scaleworms, usually covered with thick, loosely constructed tubes consisting of thin threads filled with clay or sand particles. The most important recent revision was made by Strelzov (1968b).
Key to Genera la. lb. 2a (la). 2b (la). 3a (2a). 3b (2a). 4a (2b). 4b (2b). 5a (lb). 5b (lb). Ommatophores present Ommatophores absent Two antennae present Three antennae present Ommatophores fused in the midhne Ommatophores separated from each other Superior neurosetae long and tapering, more or less hirsute Superior neurosetae short and brush-topped Three antennae present Antennae absent 2 5 3 4 Neopanthatis Eupolyodontes Polyodontes Panthalis Eupanthalis Resno
Generic Definitions Eupanthalis Mclntosh 1876, E. kinbergi Mclntosh 1876; 8 species. Ommatophores absent, three antennae present; median antenna attached dorso-posteriorly. Setae include slender, pilose tapering capillaries; bluntly acicular, aristate ones with pilose shafts, and side-shaped, dentate ones. Eupolyodontes Buchanan 1894, E. cornishii Buchanan 1894; 5 species. Ommatophores present and separated from each other. Two frontal antennae present; small nuchal papilla present. Setae include bipinnate pencillate setae; serrulate setae and smooth, acicular spines. Neopanthalis Strelzov 1968b, N. pelamida Strelzov 1968b; only species. Ommatophores present but fused in the midline. Lateral antennae short and inserted dorsally on the ommatophores. Panthalis Kinberg 1855, P. oerstedii Kinberg 1855; 12 species. Ommatophores present and separated; three antennae present. Setae include short brush-topped setae; thick, blunt acicular setae with rows of teeth subdistally and dentate sicle-shaped setae. Polyodontes Renier in Audouin and Milne Edwards 1832, Phyllodoce maxillosa Ranzani 1817; 12 species. Ommatophores present and separated; three antennae present. Setae include long, tapering hirsute ones; stout acicular setae with or without aristae, usually hirsute or dentate; and dentate sickle-shaped setae. Restio Moore 1903, R. aenus Moore 1903; only species. Ommatophores and antennae absent. Setae include long, slender ones with awnlike spines, thick, colorless and tapering, slightly curved with pilose middle part, acicular in appearance, thick, with broad lance-shaped ends and transverse rows of fine hairs.
Taxonomic Note Pseudeupanthalis Fauvel (1957), described in this family, appears to be synonymous with Sthenelanella in the family Sigalionidae.
Invalid Genera Acoetes Audouin and Milne Edwards 1832, see Polyodontes Euarche Ehlers 1887, sec Eupanthalis Eupompe Kinberg 1855, see Polyodontes Lepidio Savigny in Audouin and Milne Edwards 1832, see Panthalis.
Pholoididae NEW NAME Aphroditaceans with flattened bodies. One median antenna present. Four jaws present. Fringed elytrae alternate with dorsal cirri in all setigers. Each elytron with concentric rings. Neurosetae composite, notosetae simple. The family consists of one genus, Pholoides Pruvot 1895 with genotype P. dorsipapillata Pruvot 1895. The family-name is to replace Peisidicidae Hartman
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FIGURE 16. (A), Family PHOLOIDIDAE, Phoioides spera. Mugu Submarine Canyon, Caiifomia, about 100 m, 50x; (B), elytron of the above, 50x; (C), Family EULEPETHIDAE, Grubeuiepisfimbriata, Consag Rock, Golfo de California, elytron, 11 x; (D), diagram of tiie anterior end of the above, 8x; (E), median parapodium of the above, 25x; (F), anterior end of the above, first two elytrophores removed, lOx.
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and Fauchald 1971, based on the genus Peisidice Johnson 1897. There appears to be no reason to maintain a separation between the two genera; the older name has priority, and the family name should be based on this generic name. Four species are presently considered valid.
FAMILY EULEPETHIDAE CHAMBERLIN 1919C
Invalid Genera Parapholoe Hartmann-Schrder 1965, see Phohides Peisidice Johnson 1897, seePhoioides
Aphroditaceans with flattened bodies. Two antennae, four jaws present. Elytrae alternate with dorsal cirri in anterior setigers, present on all posterior setigers; elytrae marginally notched or with flattened marginal lappets. All setae simple. Neuracicula distally hammerheaded. The euiepethids (formerly pareulepids SENSU Hartman) were recently the subject of a monographic study by Pettibone (1969e). The present survey follows hers closely in most matters.
Key to Genera la. lb. 2a 2b 3a 3b (la). (la). (lb). (lb). Elytrae 12 pairs, increasing in length progressively, followed by posterior pseudelytrae 2 Elytrae more than 12 pairs; the first 12 pairs increasing in length posteriorly; the more posterior pairs smaller , 3 Elytrae with lateral border notched Pareulepis Elytrae with flattened lateral marginal lappets Grubeulepis Elytrae with lateral borders notched Eulepethus Elytrae with flattened lateral marginal lappets Mexieulepis
Taxonomic Note The pseudelytrae referred to in the key are sessile in contrast to the true elytrae, which are equipped with distinct elytrophores. They are in the same general position as the elytrae and appear to have similar functions. Generic Definitions Eulepethus Chamberlin 1919c, Eulepis hamifera Grube 1875; only species. Body with 60-70 segments. Elytrae with marginal notches on segments 2,4,5,7 . . . 21,24 and from segment 28 small elytrae with entire margins on every segment. Grubeulepis Pettibone 1969e, Eulepis fimbriata Treadwell 1901; 7 species. Body with about 38 segments. Elytrae with flattened marginal lappets including in all twelve pairs, on segments 2,4,5,7 . . . 21,24; pseudelytrae begin on segments 26-29. Mexieulepis Rioja 1961, Ai. elongam Rioja 1961; 2 species. Body with about 50 segments. Elytrae with flattened marginal lappets, including numerous pairs, on segments 2,4,5,7 . . . 21,24; smaller elytrae on every segment starting on segments 27 or 28.
Pareulepis Darboux 1899, Eulepis wyvillei Mclntosh 1855; 2 species. Body with about 37 segments. Twelve pairs of elytrae on segments 2,4,5,7 . . . 21,24; all with lateral border notched. Pseudelytrae begin on segment 28.
Invalid Genus Eulepis Grube 1875, see Eulepethus
FAMILY SIGALIONIDAE MALMGREN
1867
Aphroditaceans with quadrangular or flattened bodies. One to three antennae present; four jaws present. Marginally fringed elytrae alternate with dorsal cirri on anterior setigers and are present on all posterior setigers. Neurosetae composite, notosetae simple. Sigalionids are common in soft bottoms; they tend to be long-bodied and the scales are usually rather closely appended to the bodies. This, combined with the well-developed notopodial fascicles of setae, tend to give them a rather quadrangular appearance. The sigalionids are presently under revision by Pettibone, who has already published a series of important monographs (Pettibone 1969c, 1970a, 1970b, 1970e, 1971b, 1971c). The key given below must be considered provisional.
Key to Genera la. lb. One median antenna present At least two antennae present 2 3
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FIGURE
17. Family SIGALIONIDAE, Sthenolepis japnica. Bay of Nha Trang, Viet Nam, 50 m, 25x.
2a 2b 3a 3b 4a 4b 5a 5b 6a 6b 7a 7b 8a 8b 9a
(la). (la). (lb). (lb). (3b). (3b). (4a). (4a). (4b). (4b). (6a). (6a). (7a). (7a). (7b).
9b (7b). 10a 10b 1 la lib 12a 12b 13a 13b 14a 14b (6b). (6b). (10b). (10b). (1 lb). (lib). (12b). ( 12b). ( 13b). (13b).
Composite neurosetae with long, slender, articulated appendages Mayella Composite neurosetae short, smooth unidentate falcigers Pholoe Two antennae present Sigalion Three antennae present 4 Lateral antennae on the prostomium proper; all antennae with short ceratophores or short and inconspicuous 5 Lateral antennae fused to first setiger; median antenna with large median ceratophore 6 Eyes large, antennae with ceratophores Euthalenessa Eyes small, antennae without ceratophores Thalenessa Median ceratophore without auricles 7 Median ceratophore with auricles 10 Dorsal cirri absent on setiger 3; elytrae not sand-incrusted 8 Dorsal cirri present on setiger 3; elytrae sand-incrusted 9 Neurosetae composite spinigers with short, canaliculated appendages Leanira Neurosetae composite, unidentate falcigers with marginally serrated appendages Pareupholoe Elytrae small, leaving the middle of the dorsum uncovered; neurosetae with short bidentate appendages Eupkoloe Elytrae covering the dorsum, neurosetae with uni- or bidentate appendages of varying lengths Psammolyce Auricles large, auricles also present on the tentacular segment Horstileanira Auricles small, absent from tentacular segment 11 All neurosetae composite unidentate falcigers with relatively short, straight appendages . . Sthenelanella At least some neurosetae otherwise (bidentate, or with articulated, canaliculated or spinigerous appendages) 12 Ventral cirri covered medially by long papillae Willeysthenelais Ventral cirri smooth 13 Parapodial lobes and sty lodes covered with fine fimbriae Fimbriosthenelais Parapodial lobes and stylodes smooth 14 Long dorsal cirri on setiger 3 Neoleanira Dorsal cirri absent from setiger 3 15
70 15a 15b 16a 16b (14b). (14b). (15b). (15b).
NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY At least some neuropodial falcigers present All neurosetae spinigerous Dorsal tubercle present on setiger 3 Dorsal tubercle absent fron) setiger 3
Science Series 28 Sthenelais 16 Sthenolepis Ehlersileanira
Generic Definitions Ehlersileanira Pettibone 1970c, Sthenelais incisa Grube 1877; 2 species. Three antennae; auricles present. Dorsal cirrus on setiger 3 absent. All neurosetae spinigerous, with canaliculated appendages. Eupholoe Mclntosh 1885, E. philippinensis Mclntosh 1885; only species. Three antennae; auricles absent. Dorsal cirrus on setiger 3 present. Elytrae small, leaving middle of body uncovered. Neurosetae with short, bidentate appendages. Encrusted with sand, Euthalenessa Darboux 1899, Thalenessa digitata Mclntosh 1885; 8 species. Three small antennae, all on the prostomium, ceratophores present. Two pairs of large eyes. Neuropodia with large foUose lobes anteriorly. Neurosetae slender, composite with multiarticulated distally bidentate appendages. Fimbriosthenelais Pettibone 1971b, Sthenelais longipinnis Grube 1870a; 2 species. Three antennae; auricles present. No dorsal cirrus on setiger 3. All parapodial lobes and stylodes fimbriated. Neurosetae either simple and spinose or composite with distally bidentate, short, slender and articulated appendages. Horstileanira Pettibone 1970a, H. vanderspoeii Pettibone 1970a; 2 species. Three antennae, large auricles, auricles present also on the tentacular segment. Setiger 3 with prominent dorsal tubercle. Neurosetae simple and spinose and composite spinigers with long, canaliculated appendages; a few may be bidentate. Leanira Kinberg 1855, L. quatrefagesi Kinberg 1855; 11 species. Three small antennae; auricles absent. No dorsal cirrus on setiger 3. Neurosetae composite spinigers with relatively short, canaliculated appendages, Mayella Hartmann-Schrder 1959, M. articulata Hartmann-Schrder 1959; only species. One antenna, auricles absent. Neurosetae composite spinigers with articulated appendages. Neoleanira Pettibone 1970a, Sigalion tetragonum rsted 1845; 2 species. Three antennae; auricles present. Long dorsal cirri on setiger 3. Neurosetae composite spinigers with canaliculated, relatively short appendages.
Pareupholoe Hartmann-Schrder 1962a, P. fimbriatus Hartmann-Schrder 1962a; only species. Three antennae; auricles absent. No dorsal cirri on setiger 3. Neurosetae composite, unidentate falcigers with serrated cutting edge. Pholoe Johnston 1839, Aphrodita minuta Fabricius 1780; 10 species. One antenna; auricles absent, Neurosetae composite falcigers with short, unidentate appendages. Notosetae in part geniculate and strongly dentate. Psammolyce Kinberg 1855, P. flava Kinberg 1855; 26 species. Three antennae, auricles absent. Third setiger with long dorsal cirri. Neurosetae composite falcigers with uni- or bidentate appendages of varying lengths. Elytrae cover body, sand-incrusted. Sigalion Audouin and Milne Edwards 1832, S. mathildae Audouin and Milne Edwards 1832; 12 species. Two lateral antennae; auricles absent. Neurosetae simple and spinose or composite with appendages of varying lengths, multiarticulated and bidentate. Sthenelais Kinberg 1855, S. helenae Kinberg 1855; 45 species. Three antennae, auricles present. No dorsal cirrus on setiger 3'. Neurosetae include simple spinose and two kinds of falcigers, stout with short, bidentate appendages, and slenderer falcigers with articulated appendages. Sthenelanella Moore 1910, S. uniformis Moore 1910; 4 species. Three antennae; auricles present. No dorsal cirri on setiger 3, All neurosetae with short, unidentate appendages. Sthenolepis Willey 1905, Leanira japnica Mclntosh 1885; 20 species. Three antennae; auricles present. With dorsal tubercles on setiger 3. Neurosetae all composite spinigers with canaliculated appendages, Thalenessa Baird 1868, Sigalion edwardsi Kinberg 1855; 8 species. Three small antennae, all on the prostomium, ceratophores absent. Two pairs of small eyes. Neurosetae composite falcigers and a few simple spinose setae. Willeysthenelais Pettibone 1971b, Sthenelais diplocirrus Grube 1875; only species.
1976
71
Three antennae; auricles present. No dorsal cirri on setiger 3. Neurosetae simple spinose or composite falcigers with thick, short appendages or more slender with articulated appendages. Ventral cirri covered medially with long papillae. Invalid Genera Conconia Schmarda 1861, see Sthenelais Euleanira Horst 1916a, see Sthenelanella Eusigalion Augener 1918, see Thalenessa Eusthenelais Mclntosh 1876, see Sthenelais and Leanira Haswellia Darboux 1899, see Euthalenessa Lepidopleurus Claparde 1868, see Psammolyce Pelogenia Schmarda 1861, see Psammolyce
Polylepis Grube 1878, see Psammolyce Pseudeupanthalis Fauvel 1957, see Sthenelanella Superfamily Chrysopetalacea Elytrae absent; notosetae flattened and expanded paleae covering the dorsum. Prostomium not fused to the first setiger.
FAMILY CHRYSOPETALIDAE EHLERS
1864
Chrysopetalaceans with short or long bodies, usually strongly flattened. Three antennae. Notosetae in transverse rows, held erect over the dorsum or covering the back as tiles on a roof. Neurosetae composite. Chrysopetalids and palmyrids are very similar; the two families are often considered synonymous, in which case the name Palmyridae applies (cf. Day 1967).
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 aurfera, 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.
72
NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY Key lo Genera
Science Sries 28
la. lb. 2a (lb). 2b (lb). 3a (2b). 3b (2b).
Notosetae cylindrical rather than flattened, not covering dorsum Dysponetus Notosetae flattened paleae 2 Prostomium without a conspicuous caruncle, body long, consisting of many segments ... Bhawania Prostomium with a conspicuous caruncle overlapping the peristomium; body short 3 First segment with asymmetrical ventral cirri; dorsal paleae of two abruptly different kinds Paleanotus First segment with paired similar cirri; dorsal paleae of one kind only Chrysopetalum
Generic Definitions Bhawania Schmarda 1861, B. myriolepis Schmarda 1861; 9 species. Body with up to 300 segments, completely covered by paleae. Caruncle absent, prostomium retractile under a fold from the first setigers. Paleae of one or two kinds, broad and narrow; neurosetae composite falcigers with blades of varying lengths. Chrysopetalum Ehlers 1864, Palmyra debilis Grube 1855; 5 species. Body with about 40 segments, completely covered by paleae. Caruncle present. Paleae of one kind only; first segment with paired, similar ventral cirri. Dysponetus Levinsen 1879, D. pygmaeus Levinsen 1879; 4 species. Body with few segments, not covered by paleae. 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, ses Paleanotus Psectra Grube 1868a, see Bhawania Taphus Webster and Benedict 1887, see Dysponetus
FAMILY PALMYRIDAE KINBERG
Superfamily Pisionacea Elytrae absent; parapodia sub-biramous or uniramous; prostomium deeply imbedded in the first segment, or projecting freely between the first segments.
FAMILY PISIONIDAE SOUTHERN
1914
Pisionaceans with maximally two pairs of antennae on the prostomium. First segment with two pairs of tentacular cirri. Four jaws present. Setae composite and simple; dorsal and ventral cirri usually clvate. Siewing (1953), Laubier (1967b) and Stecher (1968) recently have reviewed the family in terms of the generic sub-division. Pisionura Hartman and Fauchald (1971) does not belong to this family (HartmannSchider 1975). The key below is after Laubier (1967b).
1858
Chrysopetalaceans with short, flattened bodies. One antenna present. Notosetae in rosettes on the notopodial tubercles. Neurosetae composite. The palmyrids are known for one genus, Palmyra Savigny 1818 with genotype/*, aurfera Savigny 1818 and possibly for one or two more species; most described forms are considered synonyms of P. aurfera or indeterminable. The family is limited to shallow water, usually sandy or generally hard substrates in warm water and appear to be most common in the western Pacific Ocean.
19. Family PISIONIDAE, Fisione oerstedi. Independencia Bay, Peru, shallow water, 25x.
FIGURE
1976
7i
Key to Genera la. Ib. 2a (ib). 2b (lb). 3a (2b). 3b (2b). Median unpaired antennae present Median unpaired antennae absent With two pairs of sinnilar cephalic appendages With three pairs of cephalic appendages of different structure First segment asetigerous and apodous; proboscis unarmed No apodous and asetigerous segment present; proboscis with four jaws Pisionella 2 Pisionidens 3 Anoplopisione Fisione
Generic Definitions Anoplopisione Laubier 1967b, A. minuta Laubier 1967b; only species, Pisionids with palps and two pairs of tentacular cirri present. Proboscis unarmed; first segment without parapodia and setae. Fisione Grube 1857, F. oerstedii Grube 1857; 11 species. Pisionids with palps and two pairs of tentacular cirri present. Proboscis with four jaws; all anterior segments with parapodia and setae. Pisionella Hartman 1939, P. hancocki Hartman 1939; only species. Pisionids with palps; two pairs of tentacular cirri and a median unpaired antenna present. Proboscis with four jaws; all anterior segments with parapodia and setae. Pisionidens Aiyar and Alikunhi 1943, Pisionella indica Aiyar and Alikunhi 1940; only species. Pisionids with two pairs of similar cephalic appendages. Proboscis with four jaws; all segments with parapodia and setae. Invalid Genera Fauveliella Tebble 1953, see Pisionidens Pisionella Aiyar and Alikunhi 1940, see Pisionidens Praegeria Southern 1914, see Fisione
Suborder Nereidiformia Phyllodocida with at least one pair of antennae; at least one pair of tentacular cirri; palps short and usually distally blunt, frequently biarticulated. Eversible pharynx, if armed, with one pair of lateral jaws and sometimes with accessory denticles. First parapodia lateral.
FAMILY HESIONIDAE SARS
1862
Relatively short-bodied, dorsoventrally flattened worms. Two or three antennae (antennae rarely absent); palps may be absent or have from one to three articles. Two to eight pairs of tentacular cirri present. Jaws may be present. Parapodia uniramous or biramous, but notopodia always reduced compared to the neuropodia. Dorsal cirri long and slender. Neurosetae composite; notosetae, if present, simple. The hesionids are one of the least known families of polychaetes; the generic sub-division suggested by the key below is very tentative; a large number of additional genera may be expected described within a few years. Hesionids are common animals in hard substrates and in shallow water; they are more rarely found in deep water. They tend to be fragile and fragment easily upon collection; generally hesionids should be handled as scale-wotms: each specimen preserved separately.
Key to Genera la. lb. 2a (lb). 2b (lb). 3a (2a). 3b (2a). 4a (3a). 4b (3a). 5a (4b). 5b (4b). 6a (5a). 6b (5a). 7a (2b). 7b (2b). Two pairs of tentacular cirri; five antennae At least three pairs of tentacular cirri; maximally three antennae Four or more pairs of tentacular cirri Three pairs of tentacular cirri Five or more pairs of tentacular cirri Pour pairs of tentacular cirri Five pairs of tentacular cirri Six or more pairs of tentacular cirri Seven or more pairs of tentacular cirri Six pairs of tentacular cirri Seven pairs of tentacular cirri Eight pairs of tentacular cirri Parapodia uniramous Parapodia biramous Hesiosyllis 2 3 7 4 8 Friedericielta 5 6 12 Periboea 22 Orseis Alikunhia
74 8a (3b). 8b (3b). 9a (8a). 9b (8a). 10a (8b). 10b (8b). 1 la (10b).
NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY Parapodia distinctly uniramous Parapodia sub-biramous or biramous First setiger with large hooks First setiger without hooks Antennae absent Antennae present Tentacular cirri on three segments (1-2-1); dorsal cirri articulated
Science Series 28 9 10 Stmwela Hesiocaeca Bonuania 11 Hesionella
FIGURE 20. (A), Family HESIONIDAE, Hesione interlexia, Puerto Rico, intertidal, lOx; (B), Family SYLLIDAE, TyposylHs armillaris. El Descanso, Baja California, intertidal, median paiapodium, 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, 1 Calamyzas sp., diagrammatic oudine from die dorsal side, 50x.
1976 1 lb (10b). 12a (5b). 12b (5b). 13a (12a). 13b (12a). 14a (13b). 14b (13b). 15a (14b). 15b (14b). 16a (15b). 16b ( 15b). 17a (12b). 17b (12b). 18a (17a). 18b (17a). 19a (18b). 19b (18b). 20a (19b). 20b (19b). 21a (17b). 21b (17b). 11?L (6b). 22b (6b). 23a (22a). 23b (22a). 24a (23b). 24b (23b). 25a (24b). 25b (24b). 26a (23b). 26b (23b). 27a (22b). 27b (22b). 28a (27b). 28b (27b). 29a (28b). 29b (28b). 30a (29b). 30b (29b).
75 Hesionides 13 17 Syllidia 14 "Syllidia" 15 Parasyllidea 16 Neopodarke Micropodarke 18 21 Microphthalmus 19 Heteropodarke 20 Ophiodromus "Podarke" Parahesione Nereimyra 23 27 24 26 Amphiduros 25 Leocratides Leocrates Gyptis Podarkeopsis Hesiospina 28 Hesione 29 Wesenbergia 30 Dalhousiella Kefersteinia
Tentacular cirri on two segments (2-2); dorsal cirri smooth All parapodia uniramous At least some parapodia sub-biramous or biramous First three segments dorsally reduced Maximally first segment dorsally reduced Tentacular cirri on four segments (first reduced) so that they appear as 3-2-1 Tentacular cirri on three segments (2-2-2) Pharynx distally with a circlet of fine fimbriae Pharynx with either eleven or twenty-one distal papillae Pharynx with 21 distal papillae Pharynx with eleven distal papillae Three antennae present Two antennae present Median antenna attached medially or posteriorly on the prostomium Median antenna attached frontally Palpi simple Palpi biarticulated Setae present from the second segment Setae present from the fourth segment Dorsal cirri smooth Dorsal cirri articulated Three antennae Two antennae Median antenna attached medially or posteriorly on the prostomium Median antenna attached frontally Eversible pharynx distally fimbriated Eversible pharynx distally papillated Parapodia uniramous Parapodia biramous Setae present from the second segment Setae present from the fifth (or fourth) segment Notopodia with falcate spines Parapodia uniramous Palps absent Palps present Setae present from the fourth segment Setae present from the third segment Eversible pharynx smooth Eversible pharynx distally fimbriated
Generic Definitions Alikunhia Hartman 1959, Anophthalmus erythraeus Alikunhi 1949; 4 species. Three antennae, palps and three pairs of tentacular cirri present. Parapodia biramous. Eversible pharynx distally papillated, jaws absent, Amphiduros Hartman 1958, Amphidromus setosus Hessle 1925; 3 species. Three antennae, biarticulated palps and eight pairs of tentacular cirri present (2-2-2-2). Parapodia biramous. Eversible pharynx distally fimbriated, jaws absent. Median antenna attached medially.
Bonuania Pillai 1965, B. parva. Pillai 1965; only species. Antennae absent; biarticulated palps and four pairs of tentacular cirri present. Parapodia biramous. Dalhousiella Mclntosh 1901, D. carpenteri Mclntosh 1901; 5 species. Two antennae; biarticulated palps and eight pairs of tentacular cirri present. Parapodia uniramous, first setae in third segment. Eversible pharynx distally smooth, jaws absent. Friedericiella Laubier 1967c, Hesioneila pacifica Friedrich 1956; only species.
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Science Series 28
Three antennae, simple palps and five pairs of tentacular cirri present. Parapodia biramous, first setae in fourth segment. Gyptis Marion and Bobretzky 1875, G. propinqua Marion and Bobretzky 1875; 16 species. Three antennae, biarticulated palps and eight pairs of tentacular cirri present. Parapodia biramous; first setae in second segment. Eversible pharynx with 40 distal papillae, jaws absent. Hesiocaeca Hartman 1965, H. bermudensis Hartman 1965; only species. Three antennae, biarticulated palps and four pairs of tentacular cirri present. Parapodia uniramous, first setae in third segment. Eversible pharynx with a few distal papillae, jaws absent. Hesione Savigny 1818, H. splendida Savigny 1818; 7 species. Two antennae, palps absent, eight pairs of tentacular cirri present. Parapodia uniramous, setae first in third segment. Eversible pharynx distally smooth, jaws absent. Hesionella Hartman 1939b, H. mccullochae Hartman 1939b; only species. Two antennae, palps absent, four pairs of tentacular cirri present. Parapodia sub-biramous. Hesionides Friedrich 1937, H. arenaria Friedrich 1937; 3 species. Three antennae, biarticulated palps and four pairs of tentacular cirri present. Parapodia sub-biramous to biramous, first setae in third segment. Eversible pharynx with ten distal papillae and two longer cirri, jaws absent. Hesiospina Imajima and Hartman 1964, Kefersteinia similis Kessle 1925; only species. Two antennae, biarticulated palps and eight pairs of tentacular cirri present. Parapodia biramous. Eversible pharynx with 21-27 distal papillae; jaws absent. Notopodial falcate spines present. Hesiosyllis Wesenberg-Lund 1950, H. enigmtica Wesenberg-Lund 1950; only species. Five antennae (four frontal, one dorsal), smooth palps and two pairs of tentacular cirri present. Parapodia biramous. Eversible pharynx with ten distal papillae; jaws and teeth present. Heteropodarke Hartmann-Schrder 1962a, H. heteromorpha Hartmann-Schrder 1962a; only species. Three antennae, smooth palps and six pairs of tentacular cirri present. Parapodia sub-biramous. Kefersteinia Quatrefages 1865, Psamathe cirrata Keferstein 1862; only species.
Two antennae, biarticulated palps and eight pairs of tentacular cirri present. Parapodia uniramous, setae from fourth segment. Eversible pharynx with distal circlet of fimbriae; jaws absent. Leocrates Kinberg 1866b, L. chinensis Kinberg 1866b; 11 species. Three antennae, biarticulated palps and eight pairs of tentacular cirri present. Parapodia biramous. Eversible pharynx with jaws. Median antenna attached posteriorly. Leocratides Ehlers 1908, L. filamentosa Ehlers 1908; only species. Three antennae, biarticulated palps and eight pairs of tentacular cirri present. Parapodia uniramous. Eversible pharynx with jaws. Median antenna attached posteriorly. Microphthalmus Mecznikow 1865, M. sczelkowii Mecznikow 1865; 13 species. Three antennae, simple palps and six pairs of tentacular cirri present. Parapodia sub-biramous. Eversible pharynx with distal circlet of papillae, jaws absent, Median antenna attached posteriorly. Micropodarke Okuda 1938, Kefersteinia dubia Kessle 1925; only species. Two antennae, biarticulated palps and six pairs of tentacular cirri present. Parapodia uniramous. Eversible pharynx with 11 papillae distally, jaws absent. Neopodarke Hartman 1965, N. woodsholea Hartman 1965; only species. Two antennae, biarticulated palps and six pairs of tentacular cirri present. Parapodia uniramous; first setae in fourth segment. Eversible pharynx with 21 distal papillae, jaws absent. Nereimyra Blainville 1828, Nereis punctata O.F. Mller 1776; 12 species. Two antennae, biarticulated palps and six pairs of tentacular cirri present. Parapodia sub-biramous, first setae in fourth segment. Eversible pharynx with circlet of papillae distally, jaws present. Ophiodromus Sars 1862, Nereis flexuosa delle Chiaje 1825; 11 species. Three antennae, biarticulated palps and six pairs of tentacular cirri present. Parapodia biramous, first setae in second segment. Eversible pharynx distally with many fine fimbriae; jaws absent. Orseis Ehlers 1864, O. pulla Ehlers 1864; 5 species. Three antennae, simple palps and three pairs of tentacular cirri present. Parapodia uniramous, first setae in second segment. Median antenna attached posteriorly.
1976
77 Taxonomic Notes
Parahesione Pettibone 1956, Podarke luteola Webster 1880; 2 species. Two antennae, simple palps and six pairs of tentacular cirri present. Parapodia biramous, first setae in second segment. Eversible pharynx with distal circlet of fimbriae, jaws absent. Parasyllidea Pettibone 1961, P. humesi Pettibone 1961; only species. Two antennae, biarticulated palps and six pairs of tentacular cirri present. Parapodia uniramous; first setae in third segment. Eversible pharynx with a distal circlet of fimbriae, jaws absent. First segment reduced. Periboea Ehlers 1864, P. longocirmta Ehlers 1864; only species. Two antennae, triarticulated long palps and seven pairs of tentacular cirri present. Parapodia sub-biramous, first setae in third segment. Eversible pharynx with 16-22 distal papillae, jaws absent. Podarke Ex AUCTORE; confused, possibly about 12 species. Three antennae, biarticulated palps and six pairs of tentacular cirri present. Parapodia biramous, first setae from fourth segment. Eversible pharynx with a circlet of papillae, ? jaws absent. Podarkeopsis Laubier 1961b, P. galangaui Laubier 1961b; only species. Three antennae, biarticulated palps and eight pairs of tentacular cirri present. Parapodia biramous, first setae on fourth visible segment (fifth segment). Eversible pharynx with circlet of papillae, jaws absent. Struwela Hartmann-Schrder 1959, S. noodti HartmannSchrder 1959; only species. Two antennae, biarticulated palps, four pairs of tentacular cirri present. Uniramous parapodia, first setae in third segment. First setiger with large hooks. Syllidia Quatrefages 1865, S. armata Quatrefages 1865; 5 species. Two antennae, biarticulated palps and six pairs of tentacular cirri present. Parapodia uniramous, first setae in second segment. Eversible pharynx with jaws. First to third segment dorsally reduced. Syllidia Ex AUCTORE; confused, possibly five species. Two pairs of antennae, biarticulated palps and six pairs of tentacular cirri present. Parapodia uniramous, fu-st setae in fourth segment. Wesenbergia Hartman 1955, Hesionella problemtica Wesenberg-Lund 1950; only species. Two antennae, biarticulated palps and eight pairs of tentacular cirri present, Parapodia uniramous, first setae in fourth segment.
The genus Podarke usually is considered a synonym of Ophiodromus. However, the concept as usually used, differs slightly from Ophiodromus, and has been included in the key and in the definitions. The same is true of the concept Syllidia as it has been used in the literature, compared to the version originally proposed. The key is wholly dependent on correct identification of the numbers of pairs of tentacular cirri. This is perhaps best observed on the lateral side of the animal, and with the light coming in under a low angle, so that each tentacular cirrus will cast a shadow. Most hesionids have dehiscent antennae and tentacular cirri so this may be the only means of getting the numbers of anterior appendages correctly identified. Hesionids in general are rather fragile, and have to be handled very carefully in collections; the only exceptions are the large, and rather solid-bodied members of the genus Hesione. Invalid Genera Anophthalmus Alikunhi 1949, see Alikunhia Anoplonereis Giard 1882, see Ophiodromus Castalia Savigny 1820, seeNereimyra Cirrosyllis Schmarda 1861, indeterminable Dalhousia Mclntosh 1885, see Leocra/es Halimede Rathke 1843, SeeNereimyra Mania Quatrefages 1865, see Ophiodromus Onhodromus Ehlers 1908, see Ophiodromus Oxydromus Grube 1855, seeGyptis Pseudosyllidia Czemiavsky 1882, unidentifiable Schmardiella Czemiavsky 1882, unidentifiable Stephania Claparde 1870b, see Ophiodromus Telamone Claparde 1868, see Hesione Tyrrhena Claparde 1868, seeLecra/ej
FAMILY PILARCDDAE SAINT-JOSEPH
1899
Nereidiforms with ribbon-shaped or cylindrical bodies. Two or three antennae present (rarely absent). Palps simple or biarticulated; two pairs of tentacular cirri (rarely absent). Proboscis unarmed. Parapodia biramous, but notopodia always reduced. Setae simple; notosetae sometimes as a thick spine or hook. Notosetae may be absent. Pilargiids have been the subject of two recent revisions on the generic level, Pearson (1970), and Emerson and Fauchald (1971). Pettibone (1966b) added to the generic descriptions and revised several species. Pilargiids are never really numerous, but are present in most areas in moderate numbers; they tend to be associated with shelf depths and moderately coarse to mixed sediments.
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NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY Key to Genera
Science Series 28
la. lb. 2a (la). 2b (la). 3a (2a). 3b (2a). 4a (3b), 4b (3b). 5a (4b). 5b (4b). 6a (2b). 6b (2b). 7a (6b). 7b (6b). 8a (Ib). 8b (Ib). 9a (8b). 9b (8b).
Notopodia with stout emergent hooks or spines 2 Notopodia without emergent hooks or spines 8 Notopodia with recurved emergent hooks 3 Notopodia with stout, straight spines 6 Peristomium dorsally entire Ancistargis Peristomium dorsally incised 4 Dorsal and ventral cirri reduced or absent; parapodia reduced, body subcylindrical Cabira Dorsal and ventral cirri well developed, parapodia well developed, body dorso-ventrally flattened . 5 Antennae shorter than palps; integument papillated Ancistrosyllis Antennae longer than palps, integument smooth Sigambra Prostomial antennae present; parapodia sharply set off from body Synelmis Prostomial antennae absent; parapodia distinct, but not set off from body 7 Tentacular cirri present; palps absent Litocorsa Tentacular cirri absent, palps present Parandalia Prostomial antennae and tentacular cirri absent Loandalia Prostomial antennae and tentacular cirri present 9 Prostomium with two antennae, biarticulate palps present Pilargis Prostomium with three antennae; palps without palpostyles Otopsis
Generic Definitions Ancistargis Jones 1961, A. papillosus Jones 1961; 4 species. Body flattened, with two antennae, biarticulate palps, two pairs of tentacular cirri. Peristomium dorsally entire. Antennae shorter than palps. Emergent notopodial hooks present. Ancistrosyllis Mclntosh 1879, A. groenlandica McIntosh 1879; 7 species. Body flattened, with three antennae, biarticulate palps, two pairs of tentacular cirri. Peristomium dorsally incised. Antennae shorter than palps. Emergent notopodial hooks present. Cabira Webster 1879b, C. incerta Webster 1879b; 2 species. Body cylindrical, with three antennae, biarticulate palps and two pairs of tentacular cirri. Peristomium dorsally incised. Dorsal and ventral cirri reduced, parapodia poorly developed. Emergent notopodial hooks present. Litocorsa Pearson, 1970, L. stremma Pearson 1970; only species. Body cylindrical without antennae and palps; two pairs of tentacular cirri present. Emergent notopodial spines present. Loandalia Monro 1936, L. aberrans Monro 1936; only sjwcies. Body cylindrical with biarticulate palps; antennae and tentacular cirri absent. Emergent spines absent. Otopsis Ditlevsen 1917, O. longipes Ditlevsen 1917; 3 species.
Body flattened; three antennae and two pairs of tentacular cirri present; palps present, but not articulated. Emergent spines absent. Parandalia Emerson and Pauchald 1971, P. ocularis Emerson and Pauchald 1971; 6 species. Body cylindrical; biarticulate palps present; antennae Emd tentacular cirri absent. Emergent notopodial spines present. Pilargis Saint-Joseph 1899, P. verrucosa Saint-Joseph 1899; 7 species. Body flattened; two antennae, biarticulate palps and two pairs of tentacular cirri present. Emergent spines absent. Sigambra Mller 1858, 5. grubii Mller 1858; 11 species. Body flattened, three antennae, biarticulate palps and two pairs of tentacular cirri present. Antennae longer than palps. Emergent notopodial hooks present. Synelmis Chamber in 1919c, S. simplex Chamberlin 1919c; 6 species. Body cylindrical, with three antennae, biarticulate palps and two pairs of tentacular cirri present. Emergent notopodial spines present.
Taxonomic Note Talehsapia Fauvel 1932, with genotype T. annandalei Fauvel 1932, has been considered a member of the family. As noted by Emerson and Pauchald (1971), it cannot be considered a pilargiid, and has been considered an INCESTAE SEDIS. T. annandalei as reported by
7976
79
Fauvel (1935) and Mesnil and Fauvel (1939) differ from the species as originally described; they are considered here as unidentifiable Parandalia spp. (Olga Hartman, personal communication). Invalid Genera Glyphohesione Friedrich 1951, set Synelmis Harpochaeta Korschelt 1893, s&e. Ancistrosyllis Hermundura Mller 1858, indeterminable Kynephorus Ehlers 1920, see Synelmis Phronia Webster 1879b, see Pilargis
FAMILY SYLLIDAE GRUBE
1850
Small to medium-sized nereidiform polychaetes with usually, slender bodies (sometimes dorsoventrally flattened). Three antennae and simple palps present, the latter sometimes fused to each other. Two pairs of tentacular cirri. Eversible pharynx armed with a single tooth or a circlet of smaller teeth or unarmed. Pro ventricle present in nearly all forms. Parapodia uniramous, dorsal cirri usually conspicuous, setae simple or composite. The syllids are very common shallow-water forms, and tend to be most numerous on hard substrates; however, one sub-family, Exogoninae, also is well
represented in abyssal depths, In certain genera {Trypanosyllis, Autolytus and others), the structure of the trepan, the denticles along the cutting edge of the eversible pharynx, is of great importance. The pharynx is only rarely eversed in preserved material. The examination of the trepan can be done through the body-wall, if the specimen is very small and unpigmented. Larger specimens, more than .5 mm across, or pigmented specimens will have to be dissected. Examination of the eversible pharynx cannot be dispensed with in this family, even at the generic level, as is amply demonstrated in the key below. Similarly, close examination of the structure of the setae is also necessary, making the identification of syllids a time-consuming occupation, Parapodia from anterior, median and posterior region should be mounted on a slide for setal examination under a compound microscope, and the whole specimens should be scanned for modified setae of any kind. Recent monographs include Imajima (1966a-d 1967) who, in a series of papiers revised the Japanese syllids. Gidholm has published a series of papers on the subfamily Autolytinae (e.g. Gidholm 1962) and more are expected. Hartmann-Schrder has also concentrated considerable attention on the syllids. Revision of the California fauna is under way and may be expected within a few years (Piltz, in preparation).
Key to Genera la. lb. 2a (la). 2b (la). 3a (2b). 3b (2b). 4a (3b). 4b (3b). 5a (4a). 5b (4a). 6a (4b). 6b (4b). 7a (6b). 7b (6b). 8a (7b). 8b (7b). 9a (lb). 9b (lb). 10a (9b). 10b (9b). 1 la (9a). lib (9a). 12a (lib). 12b (1 lb). Ventral cirri absent AUTOLYTINAE 2 Ventral cirri present 9 Dorsal cirri on first setiger only Procerastea Dorsal cirri on most setigers 3 All setae simple, occipital flap present Alluaudella At least some setae composite, occipital flap absent 4 Dorsal cirri cylindrical 5 Dorsal cirri clvate or folise 6 Bayonet-setae thick-shafted; segmental cUiary bands absent Proceraea Bayonet-setae thin-shafted; ciliary bands present on every segment Autolytus Eversible pharynx unarmed Phyllosyllis Eversible pharynx with teeth 7 With projecting nuchal lobes, rather than flattened nuchal epaulettes Autosyllis Nuchal organs flattened epaulettes 8 Antennae and dorsal cirri clvate Umbellisyllis Antennae cylindrical, dorsal cirri flattened Myrianida Small forms (usually less than 8 mm); palps fused for at least half their lengths . .. EXOGONINAE 11 Larger forms, palps, if fused at all, only fused basally 10 Palps fused at base; dorsal cirri smooth or irregularly wrinkled EUSYLLINAE 23 Palpi free to base or absent; dorsal cirri articulated SYLLINAE 43 Antennae, tentacular and dorsal cirri absent Exogonella At least tentacular cirri present, usually also antennae and dorsal cirri 12 Two pairs of tentacular cirri present 13 One pair of tentacular cirri present 17
-so 13a (12a). 13b ( 12a). 14a (13b). 14b (13b). 15a (14b). 15b (14b). 16a (15b). 16b (15b). 17a (12b). 17b (12b). 18a (17b). 18b (17b). 19a ( 18a). 19b (18a). 20a (18b). 20b (18b). 21a (20a). 21b (20a). 22a (2b). 22b (20b). 23a (10a). 23b (10a). 24a (23b), 24b (23b). 25a (23a). 25b (23a). 26a (25b). 26b (25b). 27a (26b). 27b (26b). 28a (27b). 28b (27b). 29a (28b). 29b (28b). 30a (24a). 30b (24a). 31a (30b). 31b (30b). 32a (31b). 32b (31b). 33a (32b). 33b (32b). 34a (33a). 34b (33a). 35a (34b). 35b (34b). 36a (33b). 36b (33b). 37a (36b). 37b (36b). 38a (37b). 38b (37b). 39a (38a). 39b (38a). 40a (38b). 40b (38b). 41a (24b).
NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY Antennae absent Three antennae present Eversible pharynx non-muscular and sinuous Eversible pharynx at least partially muscular, straight Dorsal cirri long and filiform Dorsal cirri globular Body with dorsal globular papillae in addition to the dorsal cirri Body without dorsal globular papillae One antenna present; dorsal cirri rudimentary Three antennae present; dorsal cirri well developed if sometimes short Dorsal cirri long and slender Dorsal cirri short, ovoid or pyriform Parapodia with erect, contractile dorsal lobes Parapodia without dorsal lobes Eversible pharynx unarmed Eversible pharynx with a single anterior tooth Ventral cirri fused to lower edge of parapodia Ventrial cirri free from parapodia Dorsal cirri pyriform (flask-shaped) Dorsal cirri papilliform or ovoid Eversible pharynx unarmed Eversible pharynx either with a single tooth or with a chrlet of smaller teeth Eversible pharynx with a single large tooth Eversible pharynx with a series of teeth Dorsum covered with small papillae Dorsum without papillae Pharynx with an internal valve Pharynx without an internal valve Tentacular cirri absent; dorsal cirri rudimentary Tentacular cirri present; dorsal cirri well developed Antennae and tentacular cirri very short Antennae and tentacular cirri not noticeably short Enlarged knobbed acicula present in anterior parapodia Enlarged knobbed acicula absent Notacicula present Notacicula absent Antennae and dorsal cirri absent Antennae and dorsal cirri present Three first segments fused with six pairs of tentacular cirri Maximally two pairs of tentacular cirri on one segment One pair of tentacular cirri present Two pairs of tentacular cirri present Dorsal cirri flattened Dorsal cirri cylindrical Composite setae spinigerous Composite setae falcigerous Tentacular and dorsal cirri very large, ovoid and inflated Tentacular and dorsal cirri more or less cylindrical Middorsal tooth situated posterior in the eversible pharynx Middorsal tooth situated anterior in the eversible pharynx Anterior margin of eversible pharynx denticulated Anterior margin of eversible pharynx smooth Median body-region with furcate, thick simple setae Median body-region with composite falcigers Parapodia long, palps twice as long as prostomium Parapodia short and conical; palps maximally as long as prostomium Eversible pharynx with a series of very small teeth
Science Series 28 Exogonita 14 Campesyllis 15 Brania 16 Eurysyllis Plakosyllis Spermosyllis 18 19 20 Anguillosyllis Braniella 21 22 Exogonoides Pseudexogone Sphaerosyllis Exogone 25 24 30 41 Rhopaiosyllis 26 Pharyngeovalvata 27 Fauvelia 28 Atelesyllis 29 Streptosyllis SylUdes Eudontosyllis 31 Nudisyllis 32 Irmula 33 34 36 Lamellisyltis 35 Parapionosyllis Petitia Clavisyliis 37 Opisthodonta 38 39 40 Synsyllis Eusyllis Dioplosyllis PionosylUs Parautolytus
7976 41b 42a 42b 43a 43b 44a 44b 45a (24b). (41b). (41b). (10b). (10b). (43b). (43b). (44a).
8]
45b (44a). 46a (44b). 46b (44b). 47a (46b). 47b (46b). 48a (47b). 48b (47b). 49a (45b). 49b (45b). 50a (45a). 50b (45a). 51a (50b), 51b (50b). 52a (51b). 52b (5Ib). 53a (52b). 53b (52b). 54a (53b). 54b (53b). 55a (54b). 55b (54b).
Eversible pharynx with a limited number of large teeth 42 Body short, with few segments; large nuchai epaulettes present Amblyosyllis Body longer, with numerous segments; nuchal epaulettes absent Odontosyllis Palps absent; first segment with parapodia and setae Haplosyllides Palps present, first segment without parapodia and setae; usually with tentacular cirri 44 Eversible pharynx with a single tooth, a trepan or both 45 Eversible pharynx unarmed 46 Eversible pharynx with a trepan of several teeth; sometimes with an additional larger single tooth as well 49 Eversible pharynx with a single tooth 50 Dorsum covered with small papillae Xenosyllis Dorsum smooth 47 Parapodia with long, digitate distal lobes Branchiosyllis Parapodia without distal lobes 48 Paired, posteriorly directed nuchal lappets present Parapterosyllis Nuchal lappets absent Pseudosyllides Palpi as long as prostomium; body cylindrical Geminosyllis Palpi small and conical; body flattened Trypanosyliis Middorsal tooth attached posterioriy OpisthosylUs Middorsal tooth attached anteriorly 51 Setae simple, distally furcate or subdistally bossed Haplosyltis At least some setae composite 52 Dorsal cirri in posterior region alternately long and slender or large, bulbously fusiform Parasphaerosyllis Dorsal cirri may alternate between long and short, but all are of similar thickness 53 Tentacular and dorsal cirri with very few articles; dorsal cirri absent from second segment Paratyposyllis Tentacular and dorsal cirri with at least five articles; dorsal cirri present on second segment 54 Parapodia with pseudocomposite and simple setae in addition to the composite ones Syllis Parapodia maximally with two simple setae in addition to the composite ones 55 Both composite spinigers and falcigers present Ehlersia Only composite falcigers present Typosyllis Generic Definitions EUSYLLINAE. Three antennae, palps separated to the base. Antennae and tentacular cirri very short; large occipital flap present. Eversible pharynx unarmed. Dorsal cirri emerge well above base of parapodia. Autolyms Grube 1850, Nereis prolifra O.E. Mller 1788; 110 species. AUTOLYTINAE. Three antennae, two pairs of tentacular cirri; first dorsal cirri longer than the rest. Nuchal epaulettes present. Eversible pharynx with trepan with varying number of teeth. Each segment with ciliated band. Bayonet-setae thin-shafted. Autosyllis Imajima and Hartman 1964, A. japnica Imajima and Hartman 1964; only species. AUTOLYTINAE. Three antennae, two pairs of tentacular cirri; first dorsal cirri short. Dorsal cirri clvate or folise; nuchal projecting lobes present, Branchiosyllis Ehlers 1887, B. oculata Ehlers 1887; 5 species. SYLLINAE. Palps free to base; two pairs of tentacular cirri. Parapodia with long digitate distal lobes attached pre- and postsetally. Eversible pharynx unarmed.
Alluaudella Gravier 1905c, A. madagascariensis Gravier 1905c; 2 species. AUTOLYTINAE. Three short antennae and occipital flap present. Palps completely fused; two pairs of tentacular cirri; setae all simple. Eversible pharynx unarmed. Amblyosyllis Grube 1857, A. rhomheata Grube 1857; 8 species. EUSYLLINAE. Three long antennae, long palps separated to the base. Dorsal cirri wrinkled. Paired long nuchal epaulettes present. Eversible pharynx with six or seven bi- tri- or pentacuspid teeth, Anguillosyllis Day 1963, A. capensis Day 1963; only species. EXOGONINAE. Long slender forms with three minute antennae; one pair of tentacular cirri; eversible pharynx unarmed, Parapodia with long contractile dorsal lobes. Atelesyllis Pruvot i 930, A. rubrofasciata Pruvot 1930; only species.
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Science Series 28
Brania Quatrefages 1866, Exogone pusilla Dujardin 1851; 21 species. EXOGONINAE. Two pairs of tentacular cirri; dorsal cirri long and filiform. Doisai cirri longer than, ventral cirri as long as the setal lobes. Palpi as long as prostomium. Eversible pharynx with anterior dorsal tooth. Braniella Hartman 1965, B. pupa Hartman 1965; 2 species. EXOGONINAE. Three short, ovate antennae; one pair of tentacular cirri; dorsal cirri long, slender and smooth, eversible pharynx unarmed. Composite spinigers. Campesyllis Chamberlin 1919a, C. minor Chamberlin 1919a; only species. EXOGONINAE. Three short antennae; two pairs of tentacular cirri. Eversible pharynx sinuous and nonmuscular. Clavisyllis Knox 1957, C. alternata Knox 1957; only species. EUSYLLINAE. Three antennae; two pairs of tentacular cirri; prominent nuchal epaulettes present. Eversible pharynx smooth-rimmed with single dorsal tooth. Tentacular and dorsal cirri large, ovoid and inflated. Dioplosyllis Gidholm 1962, D. cirrosa Gidholm 1962; 3 species. EUSYLLINAE. Three antennae, two pairs of tentacular cirri; nuchal ridges present or absent. Palps very large and Ungulate; parapodia long. Eversible pharynx with middorsal tooth, smooth-riramed or with a few teeth. Ehlersia Quatrefages 1865, Syllis sexoculata Ehlers 1864; 15 species. SYLLINAE. Three antennae and two pairs of tentacular cirri; all anterior appendages articulated (except palps). Eversible pharynx with middorsal tooth. Setae include composite spinigers and falcigers and in posterior setigers one or two simple setae per fascicle. Eudontosyllis Knox 1960, E. aciculata Knox 1960; only species. EUSYLLINAE. Tentacular and dorsal cirri articulated, ventral cirri folise. Eversible pharynx with middorsal tooth and denticulated margin; occipital flap present. Notacicula present. Eurysyllis Ehlers 1864, E. tuberculata Ehlers 1864; 3 species. EXOGONINAE. Body short and flattened. Three globular antennae; tentacular cirri and dorsal cirri also globular. Eversible pharynx with a trepan with 10 teeth and a middorsal tooth present. Dorsum covered with rows of globular papillae.
Eusyllis Malmgren 1867, E. blomstrandi Malmgren 1867; 27 species. EUSYLLINAE. Three antennae and two pairs of tentacular cirri. Eversible pharynx with middorsal tooth, margin denticulated. Occipital flap may be present. Setae composite falcigers. Exogone rsted 1845, E. naidina rstcd 1845; 40 species. EXOGONINAE. Three antennae; one pair of tentacular cirri. Dorsal and ventral cirri shorter than setal lobes; dorsal cirri ovoid or papilliform. Eversible pharynx with a single tooth. Exogonella Hartman 1961, E. brunnea Hartman 1961; 2 species. EXOGONINAE. Antennae, tentacular and dorsal cirri absent, Eversible pharynx with a single tooth. Exogonita Hartman and Fauchald 1971, E. oculata Hartman and Fauchald 1971; only species. EXOGONINAE. Antennae absent; two pairs of tentacular cirri present. Eversible pharynx with a single tooth. Exogonoides Day 1963, E. antennata Day 1963; only species. EXOGONINAE. Three ovoid antennae; one pair of ovoid tentacular cirri. Dorsal cirri ovoid; ventral cirri fused to par^Kidia. Eversible pharynx unarmed. Fauvelia Gravier 1900, F. martinensis Gravier 1900; only species. EUSYLLINAE. Antennae absent; tentacular cirri absent; dorsal cirri rudimentary. Eversible pharynx unarmed. Geminosyllis Imajima 1966c, Trypanosyllis (Trypanedenta) ohma Imajima and Hartman 1964; only species. SYLLINAE. Body subcylindrical; three antennae, two pairs of tentacular cirri. Eversible pharynx with trepan of ten teeth and in addition a single large tooth. All antennae and cirri slender and articulated. Haplosyllides Augener 1922, H. floridana Augener 1922; only species. SYLLINAE. Three antennae; palps absent. First segment with parapodia and setae. E>orsal cirri long, ventral cirri short, all cirri smooth. Haplosyllis Langerhans 1879, Syllis spongicola Grube 1855; 10 species. SYLLINAE. Three antennae, two pairs of tentacular cirri; all cirri articulated and slender. Eversible pharynx with single tooth. Setae simple, distally furcate or with a subdistal boss or knob.
1976
83
Irmula Ehlers 1913, /. spissipes Ehlers 1913; only species. EUSYLLINAE. Three antennae; six pairs of tentacular cirri on three fused segments. All cirri smooth. Eversible pharynx with anterior single tooth. Lamellisyllis Day 1960, L. comans Day 1960; only species. EUSYLLINAE. Flattened, small form. Three foliaceous antennae; one pair of tentacular cirri. Eversible pharynx with single tooth. Dorsal cirri flattened; ventral cirri cylindrical. Myrianida Milne Edwards 1845, Nereis pinnigera Montagu 1808; 8 species. AUTOLYTINAE. Three antennae; two pairs of tentacular cirri. All dorsal cirri flattened; antennae cylindrical. Eversible pharynx with trepan with varying numbers of teeth. Nudisyllis Knox and Cameron 1970, N. tinihekia Knox and Cameron 1970; only species. EUSYILLINAE. Antennae and dorsal cirri absent. Eversible pharynx with a single tooth; margin smoothrimmed. Odontosyllis Claparde 1863, Syllis fidgwans Audouin and Milne Edwards 1833a; 35 species, EUSYLLINAE. Three antennae; two pairs of tentacular ciiri. Occipital flap usually present. Eversible pharynx with a series (less than 20) curved teeth. Opisthodonta Langerhans 1879, O. morena Langerhans 1879; 2 species. EUSYLLEÂE. Three antennae; two pairs of tentacular cirri; all antennae and cirri smooth. Eversible pharynx with a single large, posteriorly located tooth. Some anterior parapodia with very thick acicula. Opisthosyllis Langerhans 1879, O. brimnea Langerhans 1879; 10 species. SYLLINAE. Three antennae; two pairs of tentacular cirri; occipital flap may be present. All antennae and cirri articulated. Eversible pharynx with posteriorly attached mid-dorsal tooth; anterior margin smooth. Parapionosyllis Fauvel 1923, Pionosyllis gestarts Pierantoni 1903; 9 species. EUSYLLINAE. Three antennae; one pair of tentacular cirri. Eversible pharynx with a single tooth. Composite setae spinigerous. Parapterosyllis Hartmann-Schrder 1960a, P. sexoculata Hartmann-Schrder 1960a; 2 species. SYLLINAE. Three antennae and two pairs of tentacular cirri. Piostomium with paired posteriorly directed nuchal lappets. All appendages articulated. Eversible pharynx unarmed.
ParasphaerosylUs Monro 1937b, P. indica Monro 1937b; 4 species. SYLLINAE. Three antennae; two pairs of tentacular cini. Dorsal cirri anteriorly all slender and moniliform; posteriorly alternating between slender and large, bulbously fusiform cirri. Eversible pharynx with middorsal tooth. Paratyposyllis Hartmann-Schroder 1962b, P. paurocirrata Hartmann-Schrder 1962; 2 species. SYLLINAE. Three antennae; two pairs of tentacidar cirri; eversible pharynx with single tooth. All composite setae falcigers; one or two simple setae present in each of the posterior setigers. All antennae and cirri with less than five articles; second segment without dorsal cirri. Parautolytus Ehlers 1900, P. fasciatus Ehlers 1900; 2 species. EUSYLLINAE. Three antennae; two pairs of tentacular cirri. Antennae and cirri smooth. Eversible pharynx fmely denticulated, large tooth absent. Petitia Slewing 1955, P. amphophthalma Slewing 1955; only species. EUSYLLINAE. Three antennae; one pair of tentacular cirri. Palpi biarticulate in adults. Eversible pharynx with a single tooth. Composite setae falcigerous. Pharyngeovalvata Day 1951, P. natalensis Day 1951; only species. EUSYLLINAE. Three antennae; two pairs of tentacular cirri. Occipital flap present. Pharynx with valve; teeth absent. Phyllosyllis Ehlers 1897, P. albida Ehlers 1897, only species. AUTOLYTINAE. Three antennae; first segment setose, with two large, folise cirri. Eversible pharynx without teeth. Dorsal cirri folise. Pionosyllis Malmgren 1867, P. compacta Malmgren 1867; 31 species. EUSYLLINAE. Three antennae; two pairs of tentacular cirri. Tentacular and dorsal cirri smooth and cyhndrical. Eversible pharynx with single tooth; anterior margin smooth. Plakosyllis Hartmann-Schrder 1956, P, brevipes Hartmann-Schrder 1956; 2 species. EXOGONINAE. Body short, three globular antennae; tentacular cirri and dorsal cirri also globular. Eversible pharynx with a trepan with ten teeth; a single tooth also present. Dorsal globular papillae absent. Proceraea Ehlers 1864, P. picta Ehlers 1864; 7 species. AUTOLYTINAE. Three antennae; two pairs of tentacular cirri. Palps small and ventrally located, Eversible
84
Science Series 28
pharynx with trepan with numerous teeth. Epaulettes present. Bayonet-setae thick-shafted. Ciliary, bands absent from the setigers. Procerastea Langerhans 1884, P. nematodes Langerhans 1884; 3 species. AUTOLYTINAE. Three antennae, two pairs of tentacular cirri. Dorsal cirri on first setiger only. Eversible pharynx with trepan with numerous teeth. Pseudexogone Augener 1922, P. backstromi Augener 1922; only species. EXOGONINAE. Three anteimae; one pair of tentacular cirri. Eversible pharynx unarmed. Dorsal cirri present on second segment. Pseudosyllides Augener 1927b, P. curacaoensis Augener 1927b; only species. SYLLINAE. Three antennae; two pairs of tentacular cirri; antennae and cirri strongly articulated. Eversible pharynx unarmed with a smooth margin. Rhopalosyllis Augener 1913a, R. hamuUfera Augener 1913a; only species. EUSYLLINAE. Three antennae and two pairs of tentacular cirri. Antennae and cirri smooth. Dorsum covered with long papillae. Eversible pharynx unarmed. Spermosyllis Claparde 1864, S. torulosa Claparde 1864; 3 species. EXOGONINAE. One antenna and one pair of tentacular cirri. Eversible pharynx with a single tooth. Dorsal cirri rudimentary, ventral cirri absent. Sphaerosyllis Claparde 1863, S. hystrix Claparde 1863; 28 species. EXOGONINAE. Three antennae; one pair of tentacular cirri. Dorsal cirri pyriform (flask-shap)ed), absent on second segment. Body with adhesive papillae. Strepiosyllis Webster and Benedict 1884, S. arenae Webster and Benedict 1884; 7 species. EUSYLLINAE. Three antennae and two pairs of tentacular cirri. Eversible pharynx unarmed. Large knobbed acicula present in anterior setigers. Syllides rsted 1845, 5. longocirrata rsted 1845; 13 species. EUSYLLINAE. Three antennae and two pairs of tentacular cirri. Eversible pharynx unarmed; distal margin of pharynx smooth. No enlarged setae or acicula. Syllis Savigny 1818, S. monilaris Savigny 1818; 45 species. SYLLINAE. Three antennae; two pairs of tentacular cirri, all articulated. Eversible pharynx with a single tooth. Pseudocomposite and simple setae present in addition to the composite setae in all parts of the body.
Synsyllis Verrill 1900, S. longigularis Verrill 1900; 2 species. EUSYLLINAE. Three antennae and two pairs of tentacular cirri. Eversible pharynx with mid-dorsal tooth; margin denticulated. Middle part of body with large, distally furcate simple setae. Trypanosyllis Claparde 1864, Syllis zebra Grube 1860; 26 species. SYLLINAE. Body flattened with numerous short segments. Three antennae and two pairs of tentacular cirri; all articulated. Trepan with several teeth; a single tooth may be present. Typosyllis Langerhans \S79, Syllis krohnii Ehlers 1864; 89 species. SYLLINAE. Three antennae and two pairs of tentacular cirri; all articulated. Eversible pharynx with a single tooth. Setae include uni- or bidentate falcigers in addition to one or two simple setae in each of the posterior setigers. Umbellisyllis Sars 1869, U. fasciata Sars 1869; 2 species. AUTOLYTINAE. Three clvate antennae; two pairs of tentacular cirri. Nuchal organs foliaceous. Dorsal cirri clvate. Xenosyllis Marion and Bobretzky 1875, Syllis scabra Ehlers 1864; 2 species. SYLLINAE. Three short, thick antennae; two pairs of tentacular cirri. Tentacular cirri and dorsal cirri with few moniliform or collared articles. Dorsum covered with small papillae. Eversible pharynx unarmed, Taxonomic Notes The subfamilies have been accepted strictly as defined above; as a consequence several of the genera have beeil moved from one subfamily to another. This is not considered to be of any great importance: the differences between the subfamilies, especially between the EUSYLLINAE and SYLLINAE appear to be of more practical than scientific value. The genus Irmula was originally described in the Hesionidae. It was moved to Syllidae by Day (1967). It has a very isolated position in the family due to the presence of three modified anterior segments with six pairs of tentacular cirri. However, the structure of pharynx, parapodia and setae is typically syllid, so it appears best to retain it in the Syllidae. Hesiosyllis Wesenberg-Lund 1950, described as intermediary between the syllids and the hesionids, is treated here among the latter, in that the structure of the pharynx, setae, parapodia and tentacular cirri appear to resemble members ofthat family much more than it resembles any member of the Syllidae.
1976 Invalid Genera
85
Amytis Savigny 1818, &et Proceraea Anoplosyllis Claparde 1868, see Syllides Aporosyllis Quatrefages 1865, seeSyllis Autolytides Malaquin 1893, seeAutolytus Brachysyllis Imajima and Hartman 1964, see Dioplosyllis Chaetosyllis Malmgren 1867, seeEhlersia Cirrosyllis Schmarda 1861, see Amblyosyllis and Autolytus Claparedia Quatrefages 1865, see Eusyllis (?) Crithida Gosse 1855, seeAutolytus Cystonereis Kllikerm Koch 1846, seeExogone Desmosyllis Verrill 1900, see Eusyllis Diploceraea Grube 1850, seeAutolytus Doyeria Quatrefages in Milne Edwards 1848, NOMEN
NUDUM
Eucerastes Ehlers 1864, seeMyrianida Eurymedusa Kinberg 1866b, see Trypanosyllis Exotokas Ehlers 1864, seeExogone Gattiola Johnston 1863, see Amblyosyllis Gnathosyllis Schmarda 1861, seeSyllis Gossia Quatrefages 1865, seeExogone (?) Grubea Quatrefages 1865, seeBrania Grubeosyllis Verrill 1900, see Brania Hemisyllis Verrill 1900, see Haplosyllis Hesperalia Chamberiin 1919a, lOdontosyllis Heterosyllis Claparde 1863, indeterminable loda Johnston 1840, seeSyllis sosyllis Ehlers 1864, see Typosyllis Lalage Mller 1858, NOMEU NUDVM {see Syllis) Langerhansia Czemiavsky 1882, see Ehlersia Laomedora Kinberg 1866b, indeterminable Lapithas Kinberg 1866b, indeterminable Lophosyllis Sars 1867, indeterminable Lycastis Savigny 1818, see Typosyllis Microsyllis Claparde 1863, questionably fjogo/ie Monocerina Costa 1861a, indeterminable Nereisyllis Blainville 1828, seeSyllis Nicotia Costa 1864, see Amblyosyllis Oophylax Ehlers 1864, see Exogone Paedophylax Claparde 1868, seeExogone Pagenstecheria Quatrefages 1865, see Typosyllis Parasitosyilis Potts \9\2, NOMENNVDVM Parexogone Mesnil and Caullery 1916, seeExogone Periboea Kinberg 1866b, indeterminable Photocharis Ehrenberg 1835, indeterminable Platysyltis Grube 1878, indeterminable Podonereis Blainville 1818, seeAutolytus Polybostrichus rsted 1843a, see Proceraea Polymastus Claparde 1864, see Eurysyltis Polynice Savigny in Grube 1850, questionably Autolytus Procome Ehlers 1864, see Odontosyllis
Protogrubea Czemiavsky 1881a, seeBrania PseudosyHides Czemiavsky 1882, set Amblyosyllis Pterautolytus Ehlers 1907, seeAutolytus Pterosyllis Claparde 1863, see Amblyosyllis Sacconereis Mller 1853, seeAutolytus Salvatoria Mclntosh 1885, seeBrania Sckmardia Quatrefages 1865, Siee Exogone Stephanosyllis Claparde 1864, see Proceraea Syllia Quatrefages 1865, seeExogone Sylline Grube 1860, seeAutolytus Sylline Claparde 1864, seeExogone Tetraglene Grube 1863, see Trypanosyllis Thoe Kinberg 1866b, see Typosyllis Thylaciphorus Quatrefages 1865, see Amblyosyllis Trichosyllis Schmarda 1861, seeSyllis Virchowia Langerhans 1879, see Umbellisyllis Xenosyllides Perejaslavzeva in Jakubova 1930, questionably Umbellisyllis
FAMILY CALAMYZIDAE HARTMANN-SCHRDER
1971
Body short, anteriorly and posteriorly rounded Prostomiura small and without appendages with a suck ing mouth on the ventral side. All cirri (tentacular cirri, dorsal and ventral cirri) short and digitate. Anal cirri absent. Setae composite. Eversible pharynx with stylet shaped sucking tube, otherwise without specializations The only known genus and species, Calamyzas amphictenicola Arwidsson 1932 is parasitic on the ampharetid polychaete,/Impfcicie/i gunneri (Sars 1835) from Sweden. A review can be found in HartmannSchrder 1971.
FAMILY NEREIDAE JOHNSTON
1845
Elongated, multi-segmented nereidiform polychaetes. Two, rarely one, antennae; palps biarticulated. Two or four pairs of tentacular cirri. Eversible pharynx with a pair of jaws and often accessory denticles or papillae. Parapodia nearly always biramous, usually with complex flattened lobes and cirri. Setae composite or simple, spinigerous or falcigerous. The nereids are common forms in all depths, and penetrate fteshwater and to a very limited extent, even terrestrial environments (Pflugfelder 1933). Some nereids are easy to maintain under laboratory conditions and have been used extensively for experimental research. The most popular of the nereids thus used, is Hediste diversicolor, referred to in the experimental literature incorrectly as Nereis diversicolor in most cases; in other cases, more correctly as Neanthes diversicolor. Some critical problems are associated with the recognition of species within the family, in that all species
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cannot be recognized on strictly morphological characters (Smith 1958). It is suspected that in fact a series of widespread species (Nereis pelgica, Neanthes virens, Platynereis dumerilii and others) may turn out
to be species-complexes, defined on non-morphological features. It is thus of importance that the provenance of experimental organisms be stated clearly in all pubUcations.
Key to Genera la. lb. 2a (1 b). 2b (lb). 3a (2a). Peristomium forms a large ventral collar Peristomium not ventrally enlarged Some notopodia with pectinate branchiae Branchiae absent Branchiae arise from the dorsal cirrus; all setae composite spinigers Cheilonereis 2 3 4 Dendronereis
FIGURE 21. (A), Family NEREIDAE, Aerej vexUlosa, Boiler Bay, Oregon, tntertidal, dorsal view, lOx; (B), venu-al view of the above, lOx; (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 Ai. vexillosa from Dillon Beach, California, 50x.
]J6 3b (2a). 4a (2b). 4b (2b). 5a (4b). 5b (4b). 6a (5 a). 6b (5a). 7a (5b). 7b (5b). 8a (7a). 8b (7a). 9a (7b). 9b (7b). 10a (9b). 10b (9b). 11a (10a). 1 lb ( 10a). 12a (1 lb). 12b ( 1 lb). 13a (12b). 13b (12b). 14a (10b). 14b (10b). 15a (14a). 15b (14a). 16a (14b). 16b (14b). 17a (16b). 17b (16b). 18a (17a). 18b (17a). 19a (17b). 19b (17b). 20a (15a). 20b (15a). 21a (20a). 21b (20a). 22a (20b). 22b (20b). 23a (22b). 23b (22b). 24a (23b). 24b (23b). 25a (15b). 25b (15b). 26a (25b). 26b (25b). 27a (26a). 27b (26a). 28a (27a). 28b (27a). 29a (27b), 29b (27b).
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Branchiae arise from the notopodial lobes; some composite falcigers present Dendronereides Anterior ventnim with transverse fleshy ridges Australonereis Anterior ventrum smooth * Antennae absent ^ At least one antenna present ^ Two pairs of tentacular cirri present; anterior apodous segment absent Micronereis Three pairs of tentacular cirri present; anterior apodous segment present Cryptonereis A single median antenna present 8 Two antennae present 9 Paragnaths present on the maxillary ring; parapodia biramous (except the first two) Unanereis Paragnaths absent; parapodia uniramous Dawbinia Notocirri of parapodia 5-7 broadly elytraeform Kainonereis Notocirri of parapodia 5-7 cylindrical and cirriform 10 Notopodia strongly reduced or absent 11 Median and posterior notopodia well developed, with lobes and setae 14 Notosetae present Namanereis Notosetae absent 12 Tentacular cirri articulated Lycastoides Tentacular cirri smooth 13 Notacicula present; antennae and cirri well developed Namalycastis Notacicula absent; antennae and cirri reduced Lycastopsis Eversible pharynx with either papillae or paragnaths or both, in addition to the jaws 15 Eversible pharynx with jaws, but otherwise smooth 16 Eversible pharynx with soft papillae only 20 Eversible pharynx with at least some paragnaths 25 Two pairs of tentacular cirri; apodous segment absent Micronereides Four pairs of tentacular cirri; apodous segment present 17 Dorsal cirri attached basally on the notopodial superior lobes 18 Dorsal cirri attached distally on the notopodial superior lobes 19 Notopodial homogomph falcigers present in posterior setigers Rullierinereis Notopodial homogomph falcigers absent Nicon Superior notopodial lobes long and straplike; inferior neuropodial lobe absent Steninonereis Superior notopodial lobes large and folise; inferior neuropodial lobes present Leptonereis Pharyngeal papillae at least in part in tufts 21 Pharyngeal papillae solitary 22 All setae homogomph spinigers Tylonereis Setae include also neuropodial homogomph falcigers in posterior setigers Laeonereis Ventral cirri double at least in some setigers Ceratocephale All ventral cirri simple 23 Accessory dorsal cirri on some anterior setigers; posterior dorsal cirri long and whiplike Gymnonereis Accessory dorsal cirri absent; posterior dorsal cirri not whiplike 24 Pharyngeal papillae on both rings; inferior neuropodial lobes absent; dorsal cirri distally attached ... Tylorrhynchus Pharyngeal papillae on oral ring only; inferior neuropodial lobes present; dorsal cirri basally attached Kinberginereis Eversible pharynx with both papillae and paragnaths Leonnates Papillae absent, paragnaths present 26 Paragnaths present on one pharyngeal ring only 27 Paragnaths present on both pharyngeal rings 30 Paragnaths present on the maxillary ring only 28 Paragnaths present on the oral ring only 29 Paragnaths in eight groups, all rod-shaped Solomononereis Paragnaths in patches and bands, all conical Ceratonereis Notopodial homogomph falcigers present in posterior setigers Eunereis Notopodial homogomph falcigers absent Websterinereis
88 30a (26b). 30b (26b). 31a (30a). 31b (30a). 32a (31b). 32b (31b). 33a (32b). 33b (32b). 34a (33b). 34b (33b). 35a (30b). 35b (30b). 36a (35b). 36b (35b).
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All paragnaths conical 31 Cones and in addition either transverse or pectinate paragnaths or both present on the pharynx .. 35 All setae homogomph spinigers Nectoneanthes At least some falcigers present 32 Middle and posterior neuropodia with single homogomph falcigers Hediste Homogomph falcigers, if present, in notopodial positions 33 With blunt simple falcigers in notopodia Cirronereis Simple falcigers absent, composite falcigers present or absent 34 Notopodial homogomph falcigers present in posterior setigers Nereis Notopodial homogomph falcigers absent Neanthes Paragnaths include pectinate bars and usually small patches of cones; transverse smooth bars absent Platynereis Paragnaths include transverse smooth bars, patches of cones and sometimes pectinate bars 36 Superior notopodial lobes greatly expanded in posterior setigers; pectinate bars usually present Pseudonereis Superior notopodial lobes not expanded in any setigers; pectinate bars absent Perinereis
Generic Definitions Australonereis Hartman 1954, Nereis (Leonnates) ehlersi Augener 1913a; only species. Eversible pharynx with soft papillae on the maxillary ring, oral ring bare. Four pairs of tentacular cirri; biramous parapodia. Notosetae homogomph spinigers; neurosetae homo- and heterogomph falcigers. With fleshy transverse ridges across anterior ventrum, Ceraiocephale Malmgren 1867, C. loveni Malmgren 1867; II species. Eversible pharynx with soft papillae on both rings. Four pairs of tentacular cirri; biramous parapodia. Setae include homogomph and heterogomph spinigers and heterogomph falcigers. Ventral cirri double on at lest some setigers, usually on most. Ceratonereis Kinberg 1866a, C. mirabilis Kinberg 1866a; 53 species. Eversible pharynx with paragnaths on the maxillary ring only. Four pairs of tentacular cirri; biramous parapodia. Notosetae include homogomph spinigers and falcigers; neurosetae homo- and heterogomph spinigers and heterogomph falcigers. Dorsal cirri attached basally to the superior notopodial lobe; inferior neuropodial lobe may be present. Cheilonereis Benham 1916, Nereis cyclurus Harrington 1897; 2 species. Eversible pharynx with conical paragnaths on both rings. Four pairs of tentacular cirri; biramous parapodia. Notosetae homogomph spinigers; neurosetae homogomph spinigers and heterogomph falcigers. Superior notopodial lobes large and folise with dorsal cirrus attached medially. Peristomium greatly expanded ventrally. Cirronereis Kinberg 1866a, C. gracilis Kinberg 1866a; only sf)ecies.
Eversible pharynx with conical paragnaths on both rings. Four pairs of tentacular cirri; biramous parapodia. Notosetae include homogomph spinigers and blunt simple falcigers; neurosetae heterogomph spinigers and falcigers. Cryptonereis Gibbs 1971, C. malaitae Gibbs 1971; only species. Eversible pharynx without paragnaths or papillae. Three pairs of tentacular cirri; uniramous parapodia. Neurosetae heterogomph spinigers and falcigers. Frontal antennae absent; at maturity, parapodia biramous with capillary setae. Dawbinia Benham 1950, D. aucklandica Benham 1950; only species. Eversible pharynx without paragnaths or papillae. Two pairs of tentacular cirri; parapodia uniramous. Neurosetae homogomph spinigers and heterogomph falcigers. A single median antenna present. Dendronereides Southern 1921, D. heteropoda Southem 1921; 2 species. Eversible pharynx with soft papillae on both rings. Four pairs of tentacular cirri; parapodia biramous. Branchiae present as a subdivision of the notopodial superior lobes. Neuropodial inferior lobes absent. Notosetae homogomph spinigers; neurosetae homoand heterogomph spinigers and heterogomph falcigers. Dendronereis Peters 1854, D. arborifera Peters 1854; 3 species. Eversible pharynx smooth or with soft papillae. Four pairs of tentacular cirri; parapodia biramous. Branchiae present as subdivisions of the dorsal cirri. Neuropodia in anterior setigers multifid. All setae homogomph spinigers. Eunereis Malmgren 1867, Nereis longissima Johnston 1840; 7 species.
1976
Eversible pharynx with paragnaths on oral ring only. Four pairs of tentacular cirri; parapodia biramous. Notosetae homogomph spinigers and falcigers; neurosetae homo- and heterogomph spinigers and heterogomph falcigers. Gymnonereis Horst 1919a, Gymnorhynchus sibogae Horst 1918; 2 species. Eversible pharynx with papillae on the oral ring only. Four pairs of tentacular cirri; parapodia biramous. Accessory dorsal cirri present on some anterior segments; posterior dorsal cirri long and whiplike. All setae homogomph or slightly hemigomph spinigers. Hediste Malmgren 1867, Nereis diversicolor O.F. Mller 1776; only species. Eversible pharynx with conical paragnaths on both rings. Four pairs of tentacular cirri; parapodia biramous. Notosetae homogomph spinigers. Neurosetae homoand heterogomph spinigers; heterogomph falcigers. A single homogomph falciger present in median and posterior neuropodia. Kainonereis Chamberlin 1919c, K. alata Chamberlin 1919c; only species. Eversible pharynx without paragnaths or papillae. Four pairs of tentacular cirri present; parapodia biramous. Antennae bifid at the tips; broad elytraeform dorsal cirri on parapodia 5-7. Kinberginereis Pettibone 1971a, Nereis (Leptonereis) inermis Hoagland 1920; only species. Eversible pharynx with soft papillae on the oral ring only. Four pairs of tentacular cirri; parapodia biramous. Notosetae homogomph spinigers; neurosetae homogomph and heterogomph spinigers. Inferior neuropodial lobe present; dorsal cirri basal. Laeonereis Hartman 1945, Nereis culvert Webster 1879a; 6 species. Eversible pharynx with tufts of papillae on both rings and large solitary papillae on area VI. Four pairs of tentacular cirri; parapodia biramous. Notosetae homogomph spinigers; neurosetae homogomph spinigers and falcigers, the latter in posterior setigers. Inferior neuropodial lobes present. Leonnates Kinberg 1866a, L. indicus Kinberg 1866a; 10 species. Eversible pharynx with papillae on the oral ring and paragnaths on the maxillary ring. Four pairs of tentacular cirri; parapodia biramous. Notosetae homogomph spinigers; neurosetae heterogomph falcigers with coarsely serrated blades. Leptonereis Kinberg 1866a, L. laevis Kinberg 1866a; 2 species. Eversible pharynx without papillae and paragnaths. Four pairs of tentacular cirri, parapodia biramous.
Superior notopodial lobes large and folise in posterior setigers. Notosetae homogomph spinigers; neurosetae heterogomph spinigers and falcigers, the latter with long appendages. Inferior neuropodial lobe present. Lycastoides Johnson 1903, L. alticola Johnson 1903; only species. Eversible pharynx without papillae or paragnaths. Four pairs of tentacular cirri present; parapodia uniramous. Neurosetae heterogomph falcigers and spinigers. Tentacular cirri jointed; eyes absent. Lycastopsis Augener 1922, L. beumeri Augener 1922; 6 species. Eversible pharynx without papillae or paragnaths. Three pairs of tentacular cirri present; parapodia uniramous. Neurosetae heterogomph spinigers and falcigers. Antennae and cirri reduced. Micronereides Day 1963, M. capensis Day 1963; only species. Eversible pharynx without papillae or paragnaths. Two pairs of tentacular cirri present; parapodia biramous. Apodous segment absent. All setae homogomph spinigers. Micronereis Claparde 1863, M. variegata Claparde 1863; 5 species. Eversible pharynx without papillae or paragnaths. Two pairs of tentacular cirri present; parapodia biramous. Apodous segment absent. Antennae absent. All setae homogomph spinigers. Namalycastis Hartman 1959, Paranereis abiuma Mller in Grube 1871; 18 species. Eversible pharynx without papillae or paragnaths. Four pairs of tentacular cirri; parapodia sub-biramous or uniramous. Notosetae usually absent; neurosetae heterogomph spinigers and falcigers. Neuropodia with a single setal lobe only. Notopodial superior lobes prolonged in posterior setigers. Namanereis Chamberlin 1919c, Lycastis quadraticeps Blanchard 1849; 2 species. Eversible pharynx smooth or with soft papillae. Three or four pairs of tentacular cirri present; parapodia biramous with notopodia strongly reduced. Neurosetae include heterogomph spinigers and falcigers. Neanthes Kinberg 1866a, N. vaalii Kinberg 1866a; 50 species. Eversible pharynx with conical paragnaths on both rings. Four pairs of tentacular cirri, parapodia biramous. Notosetae homogomph spinigers; neurosetae homo- and heterogomph spinigers and heterogomph falcigers. Nectoneanthes Imajima 1972, Nereis (Alitta) oxypoda Marenzeller 1879; 2 species. Eversible pharynx with conical paragnaths on both rings. Four pairs of tentacular cirri present; parapodia
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biramous. All setae homogomph spinigers. Expanded superior notopodial lobes in median and posterior seliger, with dorsal cirrus inserted between the superior and the median lobes. Nereis Linnaeus 1758, A^. pelgica Linnaeus 1758; 134 species. Eversible pharynx with conical paragnaths on both rings. Four pairs of tentacular cirri present; parapodia biramous. Notosetac include homogomph spinigers and falcigers, the latter in median and posterior setigers; neurosetae include homo- and heterogomph spinigers and heterogomph falcigers. Nicon Kinberg 1866a, N. pictus Kinberg 1866a; 15 species. Eversible pharynx without papillae or paragnaths. Four pairs of tentacular cirri; parapodia biramous. Notosetae homogomph spinigers; neurosetae homogomph spinigers and falcigers. Inferior neuropodial lobes present. Perinereis Kinberg 1866a, P. novaehollandiae Kinberg 1866a; 60 species. Eversible pharynx with conical and transverse paragnaths on both rings; four pairs of tentacular cirri; parapodia biramous. Notosetae homogomph spinigers; neurosetae homo- and heterogomph spinigers and heterogomph falcigers. Platynereis Kinberg 1866a, P. magalhaensis Kinberg 1866a; 20 species. Eversible pharynx with paragnaths on both rings, including cones, and pectinate bars. Four pairs of tentacular cirri; parapodia biramous. Notosetae homogomph spinigers and falcigers, the latter sometimes fused to form simple falcigers; neurosetae include homo- and heterogomph spinigers and heterogomph falcigers. Pseudonereis Kinberg 1866a, P. gallapagensis Kinberg 1866a; 7 species. Eversible pharynx with paragnaths on both rings, including cones, transverse smooth bars and pectinate bars. Four pairs of tentacular cirri; parapodia biramous. Notosetae include homogomph spinigers and falcigers; neurosetae homo- and heterogomph spinigers and heterogomph falcigers, Rullierinereis Pettibone 1971a, Leptonereis zebra Rullier 1963; 5 species. Eversible pharynx without papillae or paragnaths. Four pairs of tentacular cirri; parapodia biramous. Notosetae include homogomph spinigers and falcigers, the latter in posterior setigers; neurosetae homo- and heterogomph spinigers and heterogomph falcigers. Inferior neuropodial lobes present. Sotomononereis Gibbs 1971, 5. rnaranensis Gibbs 1971; only species.
Eversible pharynx with eight groups of rod-shaped paragnaths on the maxillary ring; oral ring bare. Four pairs of tentacular cirri; parapodia biramous. Notosetae include homogomph spinigers and falcigers; neurosetae hemi-, heterogomph spinigers and heterogomph falcigers. Steninonereis Wesenberg-Lund 1958, 5. martini Wesenberg-Lund 1958; only species. Eversible pharynx without papillae and paragnaths. Four pairs of tentacular cirri; parapodia biramous. Notosetae homogomph spinigers; neurosetae homoand heterogomph spinigers, and heterogomph falcigers. Superior notopodial lobes, long and straplUce in posterior setigers. Tylonereis Fauvel 1911, T. bogoyawlenskyi Fauvel 1911; 2 species. Eversible pharynx with soft papillae on the maxillary ring only. Four pairs of tentacular cirri present; parapodia biramous. All setae homogomph spinigers. Tylorrhynchus Grube 1869a, Nereis heterochaeta Quatrefages 1865; 2 species. Eversible pharynx with soft papillae on both rings. Four pairs of tentacular cirri; parapodia biramous. Notosetae hemigomph spinigers; neurosetae hemi- and heterogomph spinigers and heterogomph falcigers. Inferior neuropodial lobes absent. Unanereis Day 1962, U. macgregori Day 1962; only species. Eversible pharynx with conical paragnaths on the maxillary ring. Four pairs of tentacular cirri; parapodia biramous. Notosetae homogomph spinigers and falcigers; neurosetae homo- and heterogomph spinigers and heterogomph falcigers. A single median antenna present. Websterinereis Pettibone 1971a, Nereis tridentata Webster 1880; only species. Eversible pharynx with paragnaths on the oral ring only. Four pairs of tentacular cirri; parapodia biramous. Notosetae homogomph spinigers; neurosetae homoand heterogomph spinigers and heterogomph falcigers. Taxonomic Notes The generic subdivision of the nereids has been based mainly on the pharyngeal structures and the presence of specific kinds of setae in the parapodial rami. Both characters are subject to some variation within each genus, and especially the pharyngeal structures require acciu'ate dissection of the anterior end. Pettibone (1971a) introduced characters of the parapodial lobes (called ligules by Pettibone) as major features in the generic identification; this may be valid, but the character has the distinct drawback that it is dependent on interpretation of shapes, which is noto-
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riously dependent on the experience of the observer, and very difficult to quantify. Pettibone in the same paper lumped series of species based on overlapping ranges in different characters; this is unfortunate, since the variability within any single population of these animals has never been examined and quantified in detail. The generic key given above, reflects the added insights of Pettibone at the generic level, but the numbers of species indicated for each genus is higher than as given by Pettibone, reflecting the more conservative approach taken to lumping at the specific level in this study. Invalid Genera Aceronereis Blainville 1818, NOMEN NUDUM Alitta Kinberg in Malmgren 1865, seeNeanthes Arete Kinberg 1866a, see Perinereis Branchionereis Blainville 1818, NOMEN NUDUM Chaunorhynchus Chamberlin 1919c, seeCeratocephate Chinonereis Chamberlin 1924, see Tylorrhynchus arroceros Claparde 1863, indeterminable Cirronereis Blainville 1818, indeterminable Gnatholycastis Ehlers 1920, see Perinereis Gymnorhynchus Horst 1918, see Gymnonereis Hedyle Malmgren 1867, see Perinereis Heminereis Quatrefages 1865, indeterminable Heteronereis rsted 1843a, see Nereis phinereis Malmgren 1865, s&e Platynereis Leontis Malmgren 1867, s&e Platynereis Lepidonereis Blainville 1818, indeterminable Lipephila Malmgren 1867, êe Perinereis Lycastella Feuerbom 1932, see Namanereis Lycastis Audouin and Milne Edwards 1833b, see Ncanalycastis Lycastoides Jakubova 1930, see Namanereis Lycoris Savigny 1818, see Nereis Masligonereis Schmarda 1861, see Pseudonereis Meganereis Blainville 1818, indeterminable Naumachius Kinberg 1866a, see Pseudonereis Nectonereis Verrill 1873b, see Platynereis Nereilepas Blainville 1828, see Neanthes Nicomedes Kinberg 1866a, indeterminable Nossis Kinberg 1866a, indeterminable Paralycastis Ehlers 1920, see Perinereis Paranereis Kinberg 1866a, see Pseudonereis Phaetusa Castelnau 1842, see Hediste Phyllonereis Hansen 1882, see Pseudonereis
Pisenoe Kinberg 1866a, see Platynereis Podonereis Blainville 1826, indeterminable Praxithea Malmgren 1867, see Nereis Protolycoris Hatschek 1893, NOMEN NUDUM Stratonice Malmgren 1867, see Perinereis Tetratrocha Sveshnikov 1959, larvae, no species named Tkoosa Kinberg 1866a, see Nereis Typklonereis Hansen 1878, indeterminable Uncinereis Chamberlin 1919c, see Platynereis
FAMILY
Antonbruunidae NEW NAME
Nereidiform polychaetes with cylindrical bodies, three antennae and simple palps. Eversible pharynx unarmed. Two pairs of tentacular cirri present. Parapodia biramous, but notopodia reduced to dorsal cirri supported by internal acicula. All setae simple. Sexual dimorphism present, with small males; inquilines in bivalves. This family is known for a single genus, Antonbruunia Hartman and Boss 1965 with genotype ^4. viridis living in the mantle cavity of the bivalve Lucina fosteri Hartman and Boss.
Suborder Glyceriformia Phyllodocida with two pairs of antennae; palps and tentacular cirri absent. Proboscis either unarmed, with four jaws or with a circlet of jaws. First parapodia lateral.
FAMILY GLYCERIDAE GRUBE
1850
Glyceriform polychaetes with long, slender bodies and conical prostomia. Eversible pharynx with four jaws in a cross. Parapodia either all biramous or all uniramous. Neurosetae composite; notosetae, when present, simple. The glycerids are long, slender polychaetes with numerous segments. Perhaps their most noticeable feature is the very long, cylindrical eversible pharynx, which they will evert when disturbed. At the tip of the pharynx are four short, usually black jaws. Glycerids are present, mainly in soft sandy or muddy substrates, in all depths; they are rarely present in large numbers. They appear to be mainly carnivores.
Key to Genera la. lb. 2a (lb). 2b (lb). Parapodia uniramous throughout Parapodia biramous throughout Prostomium long, with more than three rings; aileron of jaws with lateral wing E*rostomium short, with about three rings; aileron of jaws a simple rod Hemipodus 2 Glycera Glycerella
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Prostomium long, with seven to ten vague annuli; aileron of jaws rod-shaped. Pharyngeal organs elongate oval or filamentous. Parapodia uniramous; all setae composite spinigers. Taxonomic Notes The generic subdivision, essentially in two major genera, has been stable for the last 50 years and not much change is anticipated. Identification of species, especially in certain groups of Glycera is quite difficult, and the number of species may be subject to considerable adjustment. Specific identification depends on study of the pharyngeal organs, in addition to parapodial lobes and branchiae. Phaiyngeal 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 mictoscope, 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. 22. (A) Family GLYCERIDAE, Glycera amerkam, off Santa Barbara, intertidal, lOx; (B), Glycera capitata, median parapodium, posterior view, after Hartman, 1950,64x.
FIGURE
Invalid Genera Euglycera Verrill 1881, see Glycera Hamiglycera Ehlers 1908, see Glycera Hemipodia Kinberg 1866b, seit Hemipodus Proboscidea Blainville 1825, see Glycera Rhynchobolus Claparde 1868, see Glycera Telake Chamberlin 1919c, see Glycera
FAMILY GONIADIDAE KINBERG
Generic Definitions Glycera Savigny 1818, G. unicornis Savigny 1818; 55 species. Prostomium long, with at least three, usually five to seven annuli. Aileron of jaw with lateral wing. Pharyngeal organs of tnany kinds. Notosetae simple, capillary or acicular; neurosetae composite spinigers. Glycerella Arwidsson 1899, Hemipodus magellanicus Mclntosh 1885; 2 species. Prostomium short, with maximally four annuli. Aileron of jaws rodlike. Phaiyngeal organs long and slender. Notosetae capillaries; neurosetae composite spinigers. Hemipodus Quatrefagcs 1865, Glycera rosea Blainville in Quatrefages 1865, 15 species. 1866b Glyceriforms with long and slender bodies. Prostomiiun is conical; eversible pharynx with a circlet of smaller and larger jaw-pieces. Parapodia anteriorly uniramous, posteriorly biramous, rarely all uniramous. Neurosetae composite, notosetae simple. The goniadids often are considered as part of the glycerids, but the fwo groups differ sufficiently that a recognition at the family level is warranted. They resemble the glycerids in that their most remarkable structure is the eversible pharynx, covered with pharyngeal organs, very long, usually slender and crowned with a series of teeth. The pharyngeal organs are considerably larger than in the glycerids, and are partly sclerotinized in some genera {Glycinde, Bathyglycinde).
Key to Genera la. lb. 2a (lb). All parapodia uniramous Anterior parapodia unh-amous, posterior ones biramous Eversible pharynx with organs of many kinds Progoniada 2 3
g76 2b (lb). 3a (2a). 3b (2a). 4a (2b). 4b (2b). 5a (4a). 5b (4a). 6a (4b). 6b (4b). 7a (6b). 7b (6b). 8a (7b). 8b (7b).
93 4 Bathyglycinde Glycinde 5 6 Goniada Goniadella Ophioglycera 7 Goniadides 8 Bookhoutia Goniadopsis
Eversible pharynx with organs of one or a few kinds Notosetae slender capillaries Notosetae knobbed or falcate hooded hooks Eversible pharynx with chevrons Eversible pharynx without chevrons Neuropodia with spmigers only Neuropodia with both spinigers and falcigers Neuropodia with spinigers only At least some falcigers present Notopodia reduced to a rounded knob with a few thick acicular setae and a cirrus Notopodia well developed Anterior setae all falcigers; posterior ones both spinigers and falcigers Anterior setae all falcigers, posterior ones all spinigers
Generic Definitions Bathyglycinde Fauchald 1972, B. mexicana Fauchald 1972; 3 species. Posterior segments biramous. Chevrons absent, pharyngeal organs large and of several kinds. Notosetae capillaries; neurosetae composite spinigers. Bookhoutia Mohammad 1973, B. oligognatha Mohammad 1973; only species. Posterior segments biramous. Chevrons absent; pharyngeal organs small and of one kind. Notosetae acicular; anterior neurosetae all falcigers, posterior neurosetae both falcigers and spinigers. Glycinde Mller 1858, G. multidens Mller 1858; 20 species. Posterior segments biramous. Chevrons absent; pharyngeal organs large and of several kinds. Notosetae knobbed or falcate hooded hooks; neurosetae composite spinigers. Goniada Audouin and Milne Edwards 1833b, G. emrita Audouin and Milne Edwards 1833b; 34 species. Posterior segments biramous. Chevrons present; pharyngeal organs small, mainly of one kind. Notosetae acicular or capillary; neurosetae composite spiningers. Goniadella Hartman 1950, Eone gracilis Verrill 1873b; 2 species. Posterior segments biramous. Chevrons present; pharyngeal organs of one kind. Composite spinigers and falcigers present on all neuropodia. Goniadides Hartmann-Schrder 1960a, G. aciculata Hartmann-Schrder 1960a; 2 species. Posterior segments biramous. Chevrons absent. Notopodia reduced to short rounded lobes with a cirrus and one or two coarse acicular setae. All neurosetae composite falcigers. Goniadopsis Fauvel 1928a, G. agnesiae Fauvel 1928a; 3 species.
Posterior segments biramous. Chevrons absent. Neuropodia in uniramous part of body with composite falcigers; those in biramous part, spinigers. Notosetae acicular spines.
5>
23. (A), Family GONIADIDAE, Glycinde armgera, anterior end with pharynx half everted, after Hartman, 1950, 12.5x; (B), Goniada brurmea, median parapodium, posterior view, after Hartman, 1950, 31x; (C), anterior end of the above, 25x.
FIGURE
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Ophioglycera Verrill 1885, O. gigantea Verrill 1885; 6 species. Posterior segments biramous. Chevrons absent, pharyngeal organs of one kind and short. Notosetae slender and acicular; all neurosetae composite spinigers. Progoniada Hartman 1965, P. regularis Hartman 1965; 2 species. All segments uniramous. Chevrons present. Neurosetae include composite falcigers and spinigers in all parapodia. Taxonomic Notes The genera appear well established in the Goniadidae, The only difficult point in the key above appears in dichotomy 2 on the question of a few or many kinds of pharyngeal organs. In Glycinde and Bathyglycinde at least three distinct kinds of organs are present; in most species at least two kinds are distinctly chitinized projections. In the other genera, most species have only one kind of pharyngeal organ, but in some cases two kinds may be identifiable. Invalid Genera Eone Malmgren 1866, see Glycinde Epicaste Kinberg 1866b, see Glycinde Lacharis Kinberg 1866b, indeterminable Leonnatus Kinberg 1866b, see Goniada
FAMILY LACYDONIIDAE BERGSTRM
The lacydoniids somewhat resembles the nephtyids, in that both noto- and neuropodia are developed equally, but the parapodia and setal structures, as well as the structure of the prostomium, ally them more closely with the glycerids than with any other group of polychaetes. They traditionally have been considered as an appendix to the phyllodociform polychaetes, mainly because of the lack of proboscideal armament. The lacydoniids recently were reviewed by Uschakov (1972); the present treatment follows his closely.
1914
Glyceriform polychaetes with short and slender bodies; prostomium trapezoidal in outline; eversible pharynx unarmed. First parapodia uni- or biramous, all others biramous. Neurosetae composite, notosetae simple.
24. (A), Family LACYDONIIDAE, Paralacydonia sp., San Clemente Basin, California, 1500 m, 25x; (B), median parapodium of the above, SOx.
FIGURE
Key to Genera la. lb. 2a (lb). 2b (lb). Tentacular cirri present Tentacular cirri absent Antennae short and biarticulated; first setiger uniramous Antennae long and smooth; first setiger biramous Lacydonia 2 Paralacydonia Pseudolacydonia
Generic Definitions Lacydonia Marion and Bobtetzky 1875, L. miranda Marion and Bobretzky 1875; 4 species. Antennae short and smooth; tentacular cirri present; first twcethree parapodia uniramous. Paralacydonia Fauvel 1913, P. paradoxa Fauvel 1913; 3 species. Antennae short and biarticulated; tentacular cirri absent; only first parapodia uniramous.
Pseudolacydonia Rullier 1965b, P. caeca Rullier 1965b; only species. Antennae long and slender, not articulated; tentacular cirri absent; all parapodia birjimous.
The following five families do not appear to be closely related to any other I%yllodocida nor are they obviously related to each other. They are listed in alphabetical order.
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The three families with pelagic members, lospilidae, Tomopteridae and Typhloscolecidae, usually are considered with the bulk of the other pelagic polychaetes, most of which are related to the Phyllodocidae. This connection seems based on adaptive convergencies to the pelagic environment, such as a frequently folise condition of the parapodial lobes; a reduction in the number and importance of the setae and the lightly built, often translucent bodies in these forms. The nephtyids and sphaerodorids either are considered 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 seliger carries small, but recognizable 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 fiimily within the order Phyllodocida. The sphaerodorids, however, appear to have an extensive 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 appendages), 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 suborder, it here is considered best to leave all five as separate entities, without any commitment as to further affiliation.
FAMILY IOSPILIDAE BERGSTRM
The iospilids oen are considered allied with the phyllodocids, but appear to differ rather sharply from members of this family in most of the characters usually considered at the familial level. It is here considered a member of the order Phyllodocida, but has not been assigned to any suborder or super-familial group. The family was revised by Dales and Peter (1972) and Uschakov (1972).
1914
Pelagic, small, slender Phyllodocida. Prostomium without antennae; two short palps present. Parapodia uniramous, all setae composite. Eversible pharynx present, armed in some cases.
FIGURE 25. Family ISOPILIDAE, Phalacrophorus pictus borealis, modified after Uschakov, 1972, 50x.
Key to Genera la. 1 b. 2a (lb). 2b (lb). 3a (2b). 3b (2b). Eversible pharynx with a parr of curved jaws Eversible pharynx unarmed Up to 11 anterior segments with reduced parapodia Parapodia fully developed from segment 5 Dorsal and ventral cirri present, but small on segments 2-3 Dorsal and ventral cirri absent on segments 2-3 Phalacrophorus 2 ospilopsis 3 Paraiospilus lospilus
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Maximally ten anterior segments with reduced parapodia. Eversible pharynx with paired jaws.
FAMILY NEPHTYIDAE GRUBE
lospilopsis Augener 1922, /. antillensis Augener 1922; only species. Up to U anterior segments with rudimentary parapodia; jaws absent. lospilus Viguier 1886, /. phalacroides Viguier 1886; 2 species. Maximally four anterior segments with reduced parapodia; dorsal and ventral cirri absent on segments 2 and 3; jaws absent. Paraiospilus Viguier 1911, P. affinis Viguier 1911; only species. Maximally four anterior segments with reduced parapodia; dorsal and ventral cirri present, but rudimentary on segments 2 and 3; jaws absent. Phalacrophorus Greeff 1879, P. pictus Greeff 1879; 3 species.
1850
Phyllodocida with long, slender bodies with quadrangular cross-sections. One pair of antennae and one pair of very short palps present. A pair of lateral jaws present. Parapodia biramous with both rami well developed with complex pre- and postsetal lobes. All setae simple. Interramal, respiratory cirri present in nearly all foims. The nepthyids may superficially be confused with the sigalionids in that both groups have long, straightsided bodies abruptly tapering anteriorly, and rather more gently posteriorly. However, the nephtyids lack obvious, long appendages anteriorly, and of course the scales present in the sigalionids. Nephtyids are present at all depths, and are most common in sandy and muddy substrates.
FIGURE 26. (A), Family NEPHTYIDAE, Nephtys catiforniensis, 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, Tomopteris sp., off central California, pelagic, 5x; (E), parapodium of the above, lOx.
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Key to Genera la. Ib. 2a (lb). 2b (lb). 3a (2b). 3b (2b). Interramai cirri rudimentary or absent Interramal cirri well developed, recurved or involute Interranial cirri recurved Interramal cirri involute Eversible pharynx with subterminal and terminal papillae Eversible pharynx without subterminal and terminal papillae Micronephtkys 2 Nephtys 3 Agiaophamus Inermonephtys
Generic Definitions Agiaophamus Kinberg 1866b, A. lyratus Kinberg 1866b; 45 species. Eversible pharynx with 14 (rarely 16) rows of papillae. Interramal cirri involute; acicula distally hooked in most species. Inermonephtys Fauchald 1968a, Nephtys (Agiaophamus) inermis Ehlers 1887; 3 species. Eversible pharynx without papillae, Interramal cirri involute, acicula distally hooked. Micronephtkys Friedrich 1937, Nephthys minuta Thel 1879; 5 species. Eversible pharynx with 14 rows of papillae. Interramal cirri reduced or absent; acicula blunt-tipped, but not capped. Nephtys Cuvier in Audouin and Milne Edwards 1833b, N. hombergii Savigny 1818; 50 species. Eversible phiirynx with 22 rows of papillae. Interramal cirri recurved; acicula in most forms with a distal cap. Taxonomic Notes Identification of nephtyids to genus can be tricky on small specimens; these tend to have small, nearly straight interramal cirri and one could identify these as Micronephtkys, and the corresponding adults as Nephtys. Identification to species is not difficult, but care must be taken that appropriate segments are compared with each other (or with illustrations), since the shape of the parapodia changes along the length of the body. To overcome the problem of varying body-length, comparisons should be made on distinct fractions of the bodies (i.e. fust third, second third and last third).
The first start of the interramal cirri is seen best from the lateral side, and a probe must be used to lift the notopodial cirrus so that the small, barely emerging interramal cirrus can be seen on its ventral side. Recent revisions include Hartman (1950) and Fauchald (1968a). Invalid Genera Aglaopheme Kinberg 1866b, see Agiaophamus Aonis Savigny 1822, see Nephtys Diplobranchus Quatrefages 1865, see Nephtys Pellucidaria Sveshnikov 1959, larval forms, no species named. Portelia Quatrefages 1865, set Nephtys
FAMILY SPHAERODORIDAE MALMGREN
1867
Small Phyllodocida with short and thick or long, relatively slender bodies. Two to six antennae and one pair of tentacular cirri present. Eversible pharynx unarmed. Uniramous parapodia with simple or composite setae. Dorsum with two to many rows of large spherical tubercles. The sphaerodorids have been overlooked in most collections. They turn out to be quite frequent in deepwater samples, which tend to be better treated than shallow-water ones. It is probably only a question of time before they are found in relatively large numbers also in shallow water samples. Characteristically, they are short and grub-shaped or more slender, vermiform, and cannot be confused with any other group of polychaetes, except perhaps Sphaerosyllis (Syllidae), which has similar rows of tubercles, in the latter case, the dorsal cirri, along the dorsum. The setal structures, however, are quite different. Fauchald (1974b) reviewed the group.
Key to Genera la. lb. 2a (lb). 2b (lb). 3a (2a). 3b (2a). Both dorsal and ventral surfaces smooth At least two rows of dorsal macrotubercles Macrotubercles with terminal papillae Macrotubercles distally rounded Macrotubercles in four rows; terminal papillae very short Macrotubercles in two rows; terminal papillae long Levidorum 2 3 6 Sphaerephesia 4
98 4a 4b 5a 5b 6a 6b 7a 7b 8a 8b (3b). (3b). (4b). (4b). (2b). (2b). (6a). (6a). (6b). (6b).
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Ail setae simple Sphaerodorum At least some setae composite 5 All setae composite apart from the recurved hooks in the first seliger Epkesiella Both composite and simple setae in all setigers; apart from the first, which has simple recurved hooks Ephesiopsis Macrotubercles stalked 7 Macrotubercles sessile 8 Median antenna as long as, or longer than, the lateral antennae Clavodorum Median antenna shorter than the lateral antennae Sphaerodoridium All setae composite Sphaerodoropsis All setae simple Commensodorum Generic Definitions Sphaerodoridium Liitzen 1961, Sphaerodorum claparedii Greeff 1866; only species. Stalked macrotubercles without terminal papillae. Anterior end with a short median and two pairs of longer lateral antennae. Setae composite. Sphaerodoropsis Hartman and Fauchald 1971, Sphaerodorum sphaerulifer Moore 1911; 20 species. Four or more rows of macrotubercles without terminal papillae. Anterior end with a median and two or three pairs of lateral antennae. Setae composite. Sphaerodorum rsted 1843b, Ephesia gracilis Rathke 1843; 5 species. Four rows of sessile macrotubercles with terminal papillae. Anterior end with a median and two pairs of lateral antennae. Setae simple, including in most species large recmved hooks in the first seliger. Invalid Genera Bebryce Johnston 1844, s&t Sphaerodorum Ephesia Rathke 1843, see Sphaerodorum Hypephesia Perrier 1897, s&t Ephesiella Pollicita Johnston 1845, ste Sphaerodorum Thysanoplea Schmidt 1857, questionably Sphaerodorum
FAMILY TOMOPTERIDAE GRUBE
Clavodorum Hartman and Fauchald 1971, C. atlanticum Hartman and Fauchald 1971; 5 species. Stalked macrotubercles in six or eight rows; macrotubercles without terminal papillae. Anterior end with long median antenna and two pairs of lateral antennae. Setae composite. Commensodorum Fauchald 1974b, Sphaerodoridium commensalis Liitzen 1961 ; only species. Sessile macrotubercles in four rows; macrotubercles without terminal papillae. Anterior end with a median and two pairs of lateral antennae; all anterior appendages short. Setae simple. Ephesiella Chamberlin 1919c, Sphaerodorum abyssorum Hansen 1878; 9 species. Two rows of macro- and two rows of microtubercles on the dorsum; the sessile macrotubercles with terminal papillae. Anterior end with a median and one or two pairs of lateral antennae. A large recurved hook present in the first seliger in most species; all other setae composite. Ephesiopsis Hartman and Fauchald 1971, E. guayanae Hartman and Fauchald 1971; only species. Two rows of macro- and two rows of microtubercles on the dorsum; the sessile macrotubercles with terminal papillae. Anterior end with a median and two pairs of lateral antennae. A large recurved hook in the first seliger; setae in other setigers both simple and composite. Levidorum Hartman 1967, L. scotiarum Hartman 1967; only species. All tubercles and papillae absent. Anterior end blunt; anterior appendages absent. Parapodia with two or three acicula; setae semicomposite or simple. Sphaerephesia Fauchald 1972, S. longiseta Fauchald 1972; 3 species. Four rows of sessile macrotubercles with short terminal papillae. Anterior end with a median and two or three pairs of lateral antennae. Setae composite.
1848
Pelagic, transparent, flattened Phyllodocida. Prostomium fused with the first two segments; two antennae. Eversible pharynx unarmed. First segment well developed in juveniles and reduced in adults. Second segment in adults with a pair of long tentacular cirri supported by long acicula; setae absent. Parapodia biramous with elongated bases and short, folise rami. Glands of different kinds in the parapodia. The tomopterids have been referred to the phyllodociform group of polychaeles, but differs rather sharply from these in the structure of the anterior end (Uschakov 1972) and appears best considered as a free-standing family in the order Phyllodocia. A number of genera have been described, but it appears best to consider the family as consisting of only two genera. Recent revisions include Dales and Peter (1972) and Uschakov (1972),
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Key to Genera la. lb. Tentacular cirri of second segment much longer than the body; parapodial fins restricted to the far distal part of each ramus Enapteris Tentacular cirri as long as, or barely longer than the body; parapodial fins surround the distal part of the rami Tomopieris Generic Definitions Enapteris Rosa 1908, Tomopteris euchaeta Chun 1887; only species. Parapodial rami with folise lobes along the distal margin only. Tentacular cirri much longer than body. Tomopteris Eschscholtz 1825, T. onisciformis Eschscholtz 1825; 40 species. Parapodia rami with folise lobes surrounding most of the rami. Tentacular cirri shorter than, or about as long as the body.
Invalid Genera Briaraea Quoy and Gaimard 1827, questionably Tomopteris Escholtzia Quatrefages 1865, see Tomopteris Johnstonella Gosse 1853, see Tomopteris
FAMILY TYPHLOSCOLECIDAE ULJANIN
1878
Pelagic, transparent, fusiform or torpedo-shaped Phyllodocida. Prostomium without appendages; large folise nuchal lobes present. Two pairs of tentacular cirri. Eversible pharynx unarmed, but with retortorgan. Parapodia uniramous, with large folise dorsal and ventral cirri; setae simple. The typhloscolecids are poorly studied in terms of their biology and even basic biological information is missing. Recent reviews are given by Uschakov (1972) and Dales and Peter (1972).
27. (A), Family TYPHLOSCOLECIDAE, Travisiopsis lobifera, after Uschakov, 1972, about 12.5x; (B), Travisiopsis levinseni, anterior end, cirri on left side removed, after Uschakov, 1972, 25x.
FIGURE
Key to Genera la. lb. 2a (lb). 2b (lb). Prostomium with ciliated ridges dorsally and ventrally Prostomium without ciliated ridges Median prostomial papilla on the dorsal side present Prostomium without median dorsal papilla Generic Definitions Sagitella Wagner 1872, S. kowalewskii Wagner 1872; only species. Prostomium without ciliated ridges; median antenna absent. Travisiopsis Levinsen 1885, T. lobifera Levinsen 1885; 6 species. Prostomium without ciliated ridges; median antenna present (may be indistinct). Typhloscolex Busch 1851, T. muelleri Busch 1851; 6 species. Prostomium with dorsal and ventral ciliated ridges; median antenna present. Typhloscolex 2 Travisiopsis Sagitella
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The above definitions are after Uschakov (1972: 220-225); there appears to be differences in the structure of the nuchal organs in this family, and this may bt of value as a generic character in a future revision. Invalid Genera Acicularia Langerhans 1878, see Sagitella Nuchubranchiata Treadwell 1928, see Travisiopsis Plotobia Chamberlin 1919c, see Travisiopsis ORDER AMPHINOMIDA Prostomium distinct, a caruncle present and at least one antenna. Pharynx with a muscular, rasplike eversible ventral pad; jaws absent. Parapodia distinct with branching branchiae on at least some setigers. The family Spintheridae usually is associated with the Amphinomidae (Fauvel 1923a; Hartmann-Schrder 1971). The external resemblance of the ectoparasitic spintherids to the euphrosinids first were remarked upon by Sars (1850:210) and next by Johnston (1865: 127-128). Augener (1913a:87), while recognizing
the overall external similarity to the short-bodied amphinomids, found the differences sufficient to erect a new family for the single genus. Mantn (1967:10 and 21) remarked upon the unique construction of the eversible pharynx in Spinther compared to all other polychaetes. The genus also differs sharply from the order Amphinomida in general and the erection of a separate order to contain this family seems warranted. The new order, Spintherida, is listed below (p. 103).
FAMILY AMPHINOMIDAE SAVIGNY
1818
Amphinomida with either elongate or ovate and flattened bodies. One to five antennae present; palps present. Noto- and neurosetae in tufts, notosetae protective spinous setae, at least in part. Branchiae in branching tufts. The amphinomids are common in shallow water. They generally are referred to as fire-worms, since the spines can lead to general discomfort and infections, if they break off within the inflicted cuts. The amphinomids generally are highly colored shallow-water animals, with very distinct color patterns (cf. Fauvel 1953), but are also present, in less showy editions, in deeper water.
Key to Genera la. lb. 2a (lb). 2b (lb). 3a (2a). 3b (2a). 4a (3b). 4b (3b). 5 a (4a). 5b (4a). 6a (5b). 6b (5b). 7a (4b). 7b (4b). 8a (7b). 8b (7b). 9a (8b). 9b (8b). 10a (2b). 10b (2b). 1 la (10a). 1 lb ( 10a). 12a ( 1 la). 12b (lia). 13a (lib). 13b ( 1 lb). Caruncle completely absent; neurosetae simple hooks Hipponoa Caruncle present, usually well developed (may be difficult to discern in some species); neurosetae otherwise 2 Body ovate or fusiform 3 Body elongated with parallel sides and usually abruptly tapering anteriorly and posteriorly 10 Branchiae present on all setigers Branchamphinome Some setigers (anterior or posterior) without branchiae 4 One dorsal cirrus per notopodium 5 Two dorsal cirri per notopodium 7 Caruncle with three parallel longitudinal ridges Benthoscolex Caruncle long and folded with indistinct lateral folds 6 Eyes absent, fu-st pair of branchiae larger than the following ones Bathychloeia Eyes present, first branchiae not larger than the following ones Chloeia Caruncle with high central ridge and two wide flattened lateral folds Notopygos Lateral lobes of caruncle small and hidden under the central ridge, or absent 8 Caruncle wedge-shaped, without crest and folds Sangiria Caruncle with crest and folds 9 Caruncle high, loosely plaited and rugose Chloenopsis Caruncle low, narrowly plaited with a crenulated plate Parachloeia Caruncle small and inconspicuous, stretching through maximally three segments 11 Caruncle large and conspicuous, stretching through at least three segments 14 Branchiae present on all segments from the second or third 12 Branchiae limited to the anterior part of the body 13 Caruncle broadly triangular or cordate Amphinome Caruncle narrow and elongated Pareurythoe First segment with large, anteriorly directed hooks Paramphinome First segment without hooks Linopherus
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FIGURE 28. (A), Family AMPHINOMIDAE, Eurythoe complanata, San Felipe, Golfo de California, intertidal, lOx; (B), median pajapodium 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 SPINTHERIDAE, Spinlher sp., Point Barrow, Alaska, dredged, 5x.
102 14a (10b). 14b 15a 15b 16a 16b (10b). (14b). ( 14b). ( 15b). (15b).
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Caruncle longer than wide with a large smooth, sinuous median ridge nearly covering the narrow lateral parts Eurythoe Caruncle about as long as wide or wider, median ridge, if present, narrow 15 Caruncle without distinct median ridge, with a few deep transverse folds Hermodice Caruncle with a distinct, smooth narrow ridge 16 Furcate setae absent Pherecardia Furcate setae present Pherecardites
Generic Definitions Amphinome Bruguire 1789, Aphrodita rostrata Pallas 1776; 12 species. Body long; caruncle limited to the two-three first segments, broadly triangular or cordate. Etendritically branched branchiae on all segments from the second or third. Bathychloeia Horst 1910, 5. sibogae Horst 1910; only species. Body short and ovate; caruncle large, folded with lateral folds. Bipinnate branchiae from segment 5, first pair larger than the others. Dorsal cirri single. Eyes absent. Benthoscolex Horst 1912, B. coeca Horst 1912; 2 species. Body short and ovate; caruncle with three longitudinal parallel ridges. Dendritically branched branchiae from setiger 6, especially strongly developed on posterior setigers. Dorsal cirri single, eyes absent. Branchamphinome Hartman 1967, B. antrctica Hartman 1967; only species. Body ovate; caruncle with three digitate posteriorly directed lobes. Dendritically branched branchiae present on all setigers. Dorsal cirri single, eyes present. Chloeia Savigny 1818, Aphrodita flava Pallas 1766; 19 species. Body ovate, caruncle large, with indistinct lateral folds, much longer than wide. Branchiae pennate. Dorsal cirri single, eyes present. Chloenopsis new name, Chloenea atlntica Mclntosh 1885; only species. Body ovate, caruncle high, loosely plaited and rugose. Branchiae pennate. Dorsal cirri double; eyes present. Chloenea Kinberg, 1867c is considered a synonym of Chloeia. Chloenea Mclntosh, 1885 differs generically from Kinberg's genus and needs a new name. Chloenopsis is here proposed as the generic name for the species originally named Chloenea atlntica Mclntosh, 1885. Eurythoe Kinberg 1857, E. capensis Kinberg 1857; 10 species. Body long; caruncle long with a thick, sinuous, smooth median ridge covering the narrow folded lateral
folds. Dendritically branched branchiae. Dorsal cirri single, eyes present. Hermodice Kinberg 1857, Aphrodita carunculata Pallas 1766; 4 species. Body long; caruncle about as wide as long, without a distinct median ridge, with several transverse folds. Branchiae dendritically branched, bushy in appearance. Dorsal cirri single, eyes present. Hipponoa Audouin and Milne Edwards 1830, H. gaudichaudi Audouin and Milne Edwards 1830; 3 species. Body short and ovate; caruncle absent. Branchiae dendritically branched, bushy. Dorsal cirri single, eyes present. Notosetae large, retractile curved hooks. Linopherus Quatrefages 1865, Amphinome incaruncutata Peters 1854; 14 species. Small species, but elongated bodies in most forms; caruncle small and inconspicuous, reported absent in some forms. Branchiae tufted, present on some anterior setigers only. Dorsal cirri single, eyes present or absent. Notopygos Grube 1855, N. crinita Grube 1855; 20 species. Body ovate; caruncle with high central ridge and large flattened lateral lobes. Branchiae dendritically branched. Dorsal cirri double; eyes present. Parachloeia Horst 1912, P. marmorata Horst 1912; only species. Body ovate; caruncle low, narrowly plaited with crenulated plates. Branchiae poorly developed, with a few filaments only. Dorsal cirri double, eyes present. Paramphinome Sars 1869, Hipponoe Jeffreysii Mclntosh 1868; 6 species. Small forms, but long-bodied; caruncle short, Y-shaped or elongated. Branchiae tufted, limited to anterior setigers. Dorsal cirri single; eyes present or absent. First setiger with anteriorly directed hooks. Pareurythoe Gustafson 1930, P. japnica Gustafson 1930; 9 species. Body long; caruncle small, elongated, sinuous. Branchiae dendritically branched, present on most of the body. Dorsal cirri single, eyes present.
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Pherecardia Horst 1886, Hermodice striata Kinberg 1857; 4 species. Body long; caruncle with a narrow smooth ridge bordered by wide lateral lobes with deep parallel folds on both sides. Branchiae bushy. Dorsal cirri single; eyes present. Pherecardites Horst 1912, P. parva Horst 1912; 2 species. Body long; caruncle with median axis and lateral lamellae directed posteriorly. Bushy branchiae present from the first seliger. Dorsal cirri single, eyes present. Furcate neurosetae present. Sangiria Horst 1911, S. hysirix Horst 1911; only species. Body ovate; caruncle wedge-shaped without crest and folds. Branchiae with few filaments. Dorsal cirri double; eyes absent. Invalid Genera Amphibranchus Kinberg 1867c, see Hermodice Asloegia Kinberg 1867c, see Amphinome Blenda Kinberg 1867c, see Euryihoe Chloenea Kinberg 1867c, see Chloeia Chloochaeta Kinberg 1867c, see Chloeia Colonianella Kinberg 1867c, see Amphinome
Didymobranchus Schmarda 1861, indeterminable Eucarunculatus Malaquin and Dehorne 1907, see Pherecardia Lenora Grube 1878, see Amphinome Lirione Kinberg 1867c, see Notopygos Lycaretus Kinberg 1867c, seeEuryihoe Metamphinome Treadwell 1940, see Hipponoa Pleione Savigny 1818, possibly Amphinome Pseudeurythoe Fauvel 1932, see Linopherus Rastrara Haecker 1898, larval forms Strategis Kinberg 1867c, see Chloeia Thesmia Kinberg 1867c, ê Chloeia Thetisella Baird 1870, larval forms Veleda Castelnau 1842, indeterminable Zothea Risso 1826, indeterminable
FAMILY EUPHROSINIDAE WILLIAMS
1851
Amphinomida with short and thick bodies.One pair of antennae; palps absent. Neurosetae in tufts, notosetae in transverse rows on the dorsum; branching branchiae in rows between the notosetae. Euphrosinids often are considered in the amphinomids SENSV LATU. The two families are closely related, but the euphrosinids make up a distinct, compact group of forms, and it appears best to treat them separately.
Key to Genera la. 1 b. Notosetae bifurcate, with cylindrical shafts Notosetae flattened, smooth paieae Euphrosine Palmyreuphrosyne
Generic Definitions Euphrosine Savigny 1818, E. myrtosa Savigny 1818; 40 species. Short-bodied forms with short prostomium, caruncle with three longitudinal lobes. Branchiae in transverse rows in the dorsum of each segment. Setae include capillaries, and furcate setae. Palmyreuphrosyne Fauvel 1913, P. paradoxa Fauvel 1913; 2 species. Short-bodied forms with an elongated caruncle. Pectinate branchiae in three dorsal groups per segment. Parapodia transverse ridges. Notosetae flattened smooth paieae. Taxonomic Note Palmyreuphrosyne appears, as noted by Fauvel (1913) to combine characters of the euphrosinids with those of the palmyrids. Fauvel indicates that the eversible pharynx of Palmyreuphrosyne should be ventral.
smooth and cylindrical, which would ally the genus closely to Palmyra rather than to the amphinomidlike forms with their ventral plate-muscle pharynx. As indicated by Orrhage on several occasions, structure of the pharynx may not be an overwhelmingly strong character, and it appears best to await further study of these forms to decide the question. In the meantime, Palmyreuphrosyne is maintained where it was placed by its original describer, Fauvel (1913). Invalid Genus Lophonota Costa 1841, see Euphrosine ORDER SPINTHERIDA Body ovate. Prostomium with a median antenna. Pharynx retractable and cylindrical (resembles a turbe Harlan pharynx). Notopodia represented by membranous ridges supported by simple or furcate setae. Neuropodia with composite, strongly curved hooks. Ectoparasitic on sponges.
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NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY Family Spintheridae
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As the order; the family is known for a single genus, Spinther Johnston 1845, with the genotype, S. oniscoides Johnston 1845. About 12 species presently are recognized. Invalid Genera Cryptonota Stimpson 1854, see Spinther Oniscosoma Sars 1850, see Spinther ORDER EUNICIDA Prostomium distinct, with or without appendages. Eversible pharynx ventrolateral, strongly muscular and with at least one pair of jaws. Parapodia distinct, with strongly developed neuropodia and reduced notopodia. Supetfamily Eunicea Two to five pairs of lateral jaws (maxillae) and usually one pair of lower jaws (mandibles).
FAMILY ONUPEDDAE KINBERG
1865
Eunicea with two frontal and five occipital antennae. Maxillary carriers short; third carrier absent. Maxilla I smooth and curved. Notopodia represented by the base of the branchiae and the dorsal cirri, often supported by internal acicula. Setae include composite and pseudocomposite hooks and spinigers, pectinate setae, limbate setae and subacicular hooks. Most onuphids are tubicolous, some of them carry the tube around (Hyalinoecia); others are sessile, but may be able to leave their tubes in emergencies (Schfer 1972). AU species appear to be scavenging and feed on both plant and animal debris floating past their tube openings; others may actively hunt for debris. Onuphids tend to be common at all depths, and are, next to the lumbrinerids, the family of Eunicea best represented in deep water. Recent revisions were made by Fauchald (1968b, 1972).
29. (A), Family ONUPHIDAE, Nothria elegans, off Santa Barbara, California, 20 m, 15x; (B), maxillae of the above, 25x; (C), third parapodium of the above, 35x; (D), median parapodium of the above, 25x.
FIGURE
Key to Genera la. lb. 2a (la). 2b (la). 3a (2b). 3b (2b). 4a (lb). 4b (lb). 5a (4a). 5b (4a). 6a (5b). Two or more anterior setigers with prolonged parapodia and modified setae 2 Anterior parapodia not prolonged, or only first parapodium longer than the following ones .4 Two modified anterior setigers; these with uni- or bidentate hooks and capillary setae ,. Paranorthia More than two modified anterior setigers 3 Modified parapodia with strongly curved, grapple-hook shaped setae Rhamphobrachium Modified parapodia with composite falcigers Americonuphis Branchiae in part spiralled 5 Branchiae pectinate, simple or absent 7 Tentacular cirri absent Epidiopatra Tentacular cirri present 6 Frontal antennae short and conical, all dorsal cirri digitate Diopatra
7976 6b (5b). 7a (4b). 7b (4b). 8a (7a). 8b (7a). 9a (7b). 9b (7b). 10a (9b). 10b (9b).
105 Heptaceras 8 9 Hyalinoecia Paronuphis Onuphis 10 Paradiopatra Nothria
Frontal antennae long and slender, at least some dorsal cirri folise Tentacular cirri absent Tentacular cirri present Branchiae present Branchiae absent Branchiae in part pectinate Branchiae simple or absent Branchiae absent, dorsal cirri folise in some anterior setigers Branchiae present or absent, all dorsal cirri digitate
Generic Definitions Americonuphis Fauchald 1973, Diopatra magna drews 1891; 2 species. Frontal antennae short and conical; tentacular present. Five or more anterior setigers modified composite falcigers. Branchiae pectinate; dorsal digitate. Ancirri with cirri
Paradiopatra Ehlers 1887, P. fragosa Ehlers 1887; 2 species. Frontal antennae short and conical, tentacular cirri present. Anterior setigers and setae not modified. Branchiae absent, dorsal cirri folise in some setigers. Paranorthia Moore 1903, P. brevicornuta Moore 1903; 5 species. Frontal antennae short and conical; tentacular cirri present. Two modified anterior setigers present with uni' and bidentate hooks and capillary setae. Branchiae simple and straplike or bifid, dorsal cirri digitate. Paronuphis Ehlers 1887, P. gracilis Ehlers 1887; 5 species. Frontal antennae short and conical, tentacular cirri absent. Anterior setigers and setae not modified. Branchiae absent, dorsal cirri digitate. Rhamphobrachium Ehlers 1887, R. agassizi Ehlers 1887; 10 species. Frontal antennae short and conical, tentacular cirri present. Two or more anterior setigers modified with strongly curved, grapple-hooklike setae. Branchiae pectinate, dorsal cirri digitate. Taxonomic Notes Nothria often is considered a synonym of Onuphis. This is probably correct; species of both genera resemble each other closely. The separation is retained here for practical reasons, a final decision of this question will have to await the study of larger materials than is presently available. Invalid Genera Leptoecia Chamberlin 1919c, see Paronuphis Nereitube Blainville 1828, see Hyalinoecia Northia Johnston 1865, see Nothria Trapodia Baird 1870, see Onuphis
FAMILY EUNICIDAE SAVIGNY
Diopatra Audouin and Milne Edwards 1833b, D. amboinensis Audouin and Milne Edwards 1833b; 40 species. Frontal antennae short and conical; tentacular cirri present. Anterior setigers and setae not modified. Branchiae spiralled, dorsal cirri digitate. Epidiopatra Augener 1918, E. hupferiana Augener 1918; 5 species. Frontal antennae short and conical; tentacular cirri absent. Anterior setigers and setae not modified, branchiae spiralled, dorsal cirri digitate. Heptaceras Ehlers 1868, Diopatra phyllocirrus Schmarda 1861; only species. Frontal antennae long and slender; tentacular cirri present. Anterior setigers and setae not modified. Branchiae spiralled, dorsal cirri folise in some setigers. Hyalinoecia Malmgren 1867, Nereis tubicola O.F. Mller 1788; 20 species. Frontal antennae short and conical; tentacular cirri absent. Anterior setigers and setae not modified. Branchiae present, simple and straplike in most species; dorsal cirri digitate. Nothria Malmgren 1867, Onuphis conchylega Sars 1835; 40 species. Frontal antennae short and conical; tentacular cirri present. Anterior setigers and setae not modified. Branchiae simple and straplike or absent, dorsal cirri digitate. Onuphis Audouin and Milne Edwards 1833b, O. eremita Audouin and Milne Edwards 1833b; 60 species. Frontal antennae short and concial, tentacular cirri present. Anterior setigers and setae not modified. Branchiae pectinate, dorsal cirri digitate.
1818
Eunicea with fi-om one to five occipital antennae. Maxillary carriers short; third carrier absent. Maxilla I
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smooth and curved. Notopodia represented by branchiae and dorsal cirri, sometimes supported by internal acicula. Setae include composite falcigers and spinigers, limbate setae, pectinate setae and subacicular hooks. The eunicids are among the largest of polychaetes, some Eunice aphroditois have been reported as long as two meters. Most species are associated with hard
substrates and thus with shallow water (Fauchald 1969, 1970). Generally, the eunicids are considered carnivores, but some may be scavengers or live on large detrital particles. Tube-building is known for some species; others are burrowing into limestone or other calcium carbonate substrates. Major revisions include Hartman (1944a) and Fauchald (1970).
Key to Genera la. lb. 2a (la). 2b (la). 3a (2a). 3b (2a), 4a (3b). 4b (3b). 5a (2b). 5b (2b). 6a (lb). 6b (lb). Five occipital antennae present One to three occipital antennae present Tentacular cirri present Tentacular cirri absent Subacicular hooks absent Subacicular hooks present Composite setae falcigers Composite setae spinigers Branchiae present Branchiae absent One occipital antenna present Three occipital antennae present 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. aculeaia Wesenberg-Lund 1949; only species. Five occipital antennae; tentacular cirri present; branchiae present. Setae include limbate setae, pectinate setae, composite spinigers and subacicular hooks. Lysidice Savigny 1818, L. ninetta Audouin and Milne Edwards 1833a; 10 species. Three occipital antennae; tentacular cirri and branchiae absent. Setae include limbate setae, pectinate setae, composite falcigers and subacicular hooks. Marphysa Quatrefages 1865, Nereis sangunea Montagu 1815; 50 species. Five occipital antennae; tentacular cirri absent, branchiae present. Setae include limbate setae, pectinate setae, composite spinigers and falcigers and subacicular hooks. Nematonereis Schmarda 1861, A^. unicornia Schmarda 1861; 2 species. One occipital antenna; tentacular cirri and branchiae absent. Setae include limbate and pectinate setae, composite spinigers and subacicular hooks. Palola Gray 1847, Eunice siciliensis Grube 1840; 4 species. 30. (A), Family EUNICIDAE, Eunice antennata, Point Loma, California, intertidal, lOx; (B), subacicular hook of the above, lOOx; (C), median parapodium of the above, 25x.
FIGURE
2 6 3 5 Palola 4 Eunice Euniphysa Marphysa Paramarphysa Nematonereis Lysidice
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Five occipital antennae; tentacular cirri and branchiae present. Setae include limbate and pectinate setae and composite falcigers. Paramarphysa Ehlers 1887, P. longula Ehlers 1887; 4 species. Five occipital antennae; tentacular cirri and branchiae absent. Setae include limbate and pectinate setae, composite falcigers and subacicular hooks.
Nauphanta Kinberg 1865, ste Marphysa Nausicaa Kinberg 1865, set Marphysa Nereidice Blainviile 1828, sesLysidice Nereidonta Blainviile 1828, see Eunice, Marphysa and Palola Palpiglossus Wagner 1885, indeterminable Pseudopalolo Friedlnder in Woodworth 1907, see Lysidice Tibiana Lamarck 1816, ^& Eunice
Taxonomic Notes Palola frequently is considered a subgenus of, or synonymous with Eunice. The latter is a large genus, and anything that can be done to subdivide it, appears to be of value. Members of Palola differ consistently and uniquely from the rest of the family not only in the characters listed above, but also in the deep scoopshape of the mandibles. It thus appears valuable to retain the distinction, not least since it appears related to the ecology of the contained species, Paramarphysa and Marphysa are very similar and can be separated only as indicated in the key; the separation is maintained here provisionally.
FAMILY LUMBRINERIDAE MALMGREN
1867
Invalid Genera Amphelothrix Chamberlin 1919c, error for Aphelothrix, see Marphysa Amphiro Kinberg 1865, see Marphysa Amphiron Chamberlin 1919c, error for Amphiro, see Marphysa ^p/ie/oi/irix Chamberlin 1919c, see Marphysa Blainvillea Quatrefages 1865, see Nematonereis Eriphyle Kinberg 1865, set Eunice Eunice Rafinesque 1815, NOMEN NUDUM {Eunice Cuvier 1817 is the valid name for the main genus in this imvAy,fide Fauvel 1918:338). Heteromarphysa Verrill 1900, indeterminable Leodice Savigny 1818, see Eunice Lithognatha Stewart 1811, see Palola Lycidice Williams 1851, error foT Lysidice Macduffia Mclntosh 1885, see Marphysa Mayeria Verrill 1900, see Eunice
Eunicea without prostomial appendages, but sometimes with one to three nuchal papillae emerging from a pocket between the pro- and peristomium. Maxillary carriers short, third carrier absent. Maxilla I smooth and curved. Notopodia absent or represented by small, button-shaped projections, sometimes with internal acicula (but see Kuwaita below). Setae include limbate setae, simple and composite hooks. Subacicular hooks and pectinate setae are absent. Most lumbrinerids are free-living, buirowing forms in sand or mud or between algal hold-fasts and plantroots. They are among the most common polychaetes in sandy and muddy bottoms in shelf-depths, but also occur in numbers in deep-water areas. Identification of lumbrinerids is dependent on very accurate work and is not altogether easy. Most dependable appears to be the structure of the anterior setigers, the structure of the jawapparatus and the detailed structure of the hooks. Length and shape of the parapodial lobes also are valuable characters, but care must be taken in noting the exact location of the parapodium examined in relation to the total length of the animal. This often is difficult since the lumbrinerids fragment readily. An important character in most keys to species concerns the length of the posterior parapodial lobes in relation to the length of the anterior ones; usually stated as "prolonged posterior lobes," The key feature in this character, is that the posterior lobes have to be distinctly longer than the anterior ones. With a fragmented specimen this may be impossible to observe; in such cases, it is of the utmost importance that both paths in a key be explored; in other words, that no assumption is made about the structure of the posterior lobes.
Key to Gertera la. lb. 2a (lb). 2b (lb). 3a (2b), 3b (2b), Parapodia with distinct dorsal cirri Dorsal cirri absent Pharyngeal apparatus absent Pharyngeal apparatus present Branchiae absent Branchiae present Kuwaita 2 Ophiwicola 3 Lumbrineris Ninoe
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31. (A), Family IPHITIMIDAE, Iphitime loxorhynchi, from Loxorhynchus grandis, Santa Catalina Island, California, 26x; (B), Family LUMBRINERIDAE, Lumbrineris californiensis, off Point Finnin, 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 tricolor. Dillon Beach, California, intertidal, 16x; (H), fifth parapodium of the above, 95x; (I), Family LYSARETDDAE, Oenone fulgida. Bahia Magdalena, Baja California, 30 m, 26x; (J), fifteenth parapodium of the above, 52x.
FIGURE
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Key to Genera Vie Orensanz (1974a) la. lb. 2a (la). 2b (la). 3a (lb). 3b (lb). 4a (3b). 4b (3b). Hooded hooks distally bidentate 2 Hooded hooks, if present, distally multidentate (anterior parapodia may have incompletely formed hooks) 3 Maxilla IV with a series of articulated spines; mandibles posteriorly bifid; maxillary carriers large and posteriorly expanded Lumbrineriopsis Maxilla IV without denticles or articulated spines; mandibles posteriorly entire; maxillary carriers robust, subtriangular and not expanded posteriorly Lumbrinerides Digitiform branchiae emerging from the pwstsetai lobes on a number of anterior setigers; maxillae IV, or III and IV usually with denticulated cutting edge Ninoe If branchial structures present, they are in a different position; maxillae III and IV either smooth or with a few large teeth 4 Anterior parapodia with composite hooded hooks; maxilla II with three large teeth; maxilla IV greatly expanded with a central thin area Augeneria Paraf)odia with or without composite setae. Maxillae different Lumbrineris
Generic Definitions Kuwaita Mohammad 1973, K. magna Mohammad 1973; only species. Lumbrinerids with three nuchal papillae; dorsal cirri present and elongate in posterior segments; branchiae absent. Jaw apparatus present. Lumbrineris Blainville 1828, L. latreilli Audouin and Milne Edwards 1834; 158 species. Lumbrinerids without or with one to three nuchal papillae; dorsal cirri and branchiae absent. Jaw apparatus present. Ninoe Kinberg 1865, N. chilensis Kinberg 1865; 28 species. Lumbrinerids without nuchal papillae, dorsal cirri absent; branchiae present. Jaw apparatus present. Ophiuricola Ludwig 1905, O. cynips Ludwig 1905; only species. Lumbrinerids without nuchal papillae, dorsal cirri or jaw apparatus. Taxonomic Notes Orensanz (1974a) described the two genera Lumbrineriopsis and Lumbrinerides with L. mucronata (Ehlers 1908) and i. gesae Orensanz 1974a as type species respectively. Both genera may be valid. The change in concept of the previously described Augeneria and Ninoe indicated in the alternative key above appears unfortunate and is not recommended for future adoption. Admittedly the genus Lumbrineris as presently accepted, contains a large number of rather different fonns, but experience has taught me that surveys based on descriptions from the literature are considerably less valuable than ones based on actual specimens. It is now time to study the variability of the characters
we are using to define the many species, so that we may use knowledge of this variability when we attempt to describe more genera in this family. The genus Kuwaita is isolated in the family and may belong to a different, perhaps undescribed family.
Invalid Genera Aotearia Benham 1927, see Lumbrineris Augeneria Mon'o 1930, see Lumbrineris (but see above) Cenogenus Chamberlin 1919c, see Lumbrineris Eranno Kinberg 1865, set Lumbrineris Lumbriconereis Grube 1840, see Lumbrineris Scoletoma Blainville 1828, see Lumbrineris Unciniseta Bidenkap 1907, i^t Lumbrineris Zygolobus Grube 1863, see Lumbrineris Zygophyllus Grube 1863, error for Zygolobus, see Lumbrineris 1970
FAMILY IPHITIMIDAE FAUCHALD
Eunicea with a single pair of frontal antennae. One pair of maxillary carriers fused to maxilla I, which is smooth and curved; third carrier is absent. Notopodia represented by simple or branched branchiae. Setae include simple and composite falcigers. A single genus; Iphitime Marenzeller 1902, with genotype /. doederleini Marenzeller 1902, and five species are known. All Iphitimidae are inquilines in branchial chambers of crustaceans; their feeding modes are unknown; it is possible that they parasitize the crustacean host-, but little damage seems to be present on the branchial tissue of the host. Pilger (1972) and Fauchald (1970) reviewed the family.
no
NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY Invalid Genera
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Coelobranchus Izuka 1912, &ee Iphitime Enonella Stimpson 1854, indeterminable

FAMILY ARABELLIDAE HARTMAN
1944a
Eunicea without prostomial appendages. Maxillary carriers long and narrow, a third carrier present. Maxilla I smooth or basally dentate; notopodia absent. Setae include limbate setae and in a number of cases, thick, emergent spines.
Arabellids resemble lumbrinerids closely in overall body-construction, but differ sharply from the latter in that they have long narrow maxillary carriers and have three, rather than two carriers. Arabellids are never tubicolous; they may be parasitic in other animals, usually in other polychaetes, but also in echiurans. They may be parasitic as juveniles and become free-living as adults (Pettibone 1957c), or may stay parasitic throughout life (Wirn 1886). Major revisions: Hartman 1944a; Pettibone 1957c; Fauchald 1970.
Key to Genera la. 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). Acicular spines present Acicular spines absent Maxillary apparatus represented by a single rod Four or five pairs of jaws present Maxilla I distally falcate Maxilla I distally dentate Maxillary apparatus absent Maxillary apparatus present, but often reduced Five pairs of maxillae present Two or three pairs of maxillae present Mandibles fused and horseshoe-shaped Mandibles two triangular plates Maxillary carriers anteriorly bilobed and fused along most of their lengths Maxillary carriers anteriorly rounded and fused along their whole length 2 4 Drilognathus 3 Drilonereis Notocirrus Biborin 5 Arabella 6 Oligognathus 7 Labrorostratus Haematocleptes
Generic Definitions Arabella Grube 1850, Nereis iricolor Montagu 1804; 20 species. Arabellids with five pairs of maxillae and mandibles present. Parapodia without acicular spines. Biborin Chamberlin 1919a, B. ecbola Chamberlin 1919a; only species, Arabellids without maxillary apparatus, mandibles present. Parapodia without acicular spines. Drilognathus Day 1960, D. capensis Day 1960; only species. Arabellids with the maxillary apparatus reduced to a single rod. Parapodia with acicular spines. Drilonereis Claparde 1870b, Lumbriconereis filum Claparde 1868; 40 species. Arabellids with four or five pairs of maxillae; mandibles usually present. Parapodia with acicular spines; maxilla I distally falcate. Haematocleptes Wirn 1886, H. lerebellides Wirn 1886; 2 species. Arabellids with the maxillary apparatus consisting of two pairs of small plates, mandibles present and separate; unpaired carrier present. Acicular spines absent.
Labrorostratus Saint-Joseph 1888, L. parasiticus Saint-Joseph 1888, only species. Arabellids with two pairs of minute maxillae and the carriers fused for part of their length. Acicular spines absent. Notocirrus Schmarda 1861, N. chilensis Schmarda 1861; 10 species. Arabellids with four or five pairs of maxillae, mandibles present. Maxillae I dentate to the tip. Acicular spines present. Notopsilus Ehlers 1868, Lais acutus Kinberg 1865; only species. Arabellids with five pairs of maxillae and mandibles present. Acicular spines absent. Maxillae I proximally dentate. Oligognathus Spengel 1882, O. bonelliae Spengel 1882; 2 species, Arabellids with two or three pairs of maxillae present; mandibles fused into horseshoe-shaped piece. Acicular spines absent, Taxonomic Notes The generic sub-division is presently inconsistent in that in the group of arabellids with emergent acicular spines, forms with maxilla I distally falcate are con-
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sidered generically distinct from forms with maxilla I distally dentate. The corresponding separation should split the gtnus Arabella so that the forms with maxilla 1 dentate should go to the genus Nowpsilus, which has been defined in the generic definitions, but not considered in the key. This genus has been largely disregarded; it is not clear whether the best procedure would be to revise Arabella to separate the two sets of forms, or to fuse the well-known genera Notocirrus and Drilonereis to ensure generic conformity within the family. Invalid Genera Arabes Ehlers 1920, see Drilonereis Aracoda Schmarda 1861, see Arabella, Notocirrus and Lumbrineris Cenothrix Chamberlin 1919c, êe Arabella Labidognathus Caullery 1914a, sec Drilonereis
Lais Kinberg 1867, see Notopsilus Laranda Kinberg 1865, indeterminable Maclovia Grube 1871b, see Arabella Pterothrix Chamberlin 1919c, &ee Notocirrus
FAMILY LYSARETIDAE KINBERG
1865
Eunicea with one or three occipital antennae. Maxillary carriers long and narrow, a third carrier is present; maxilla I proximally smooth or dentate. Notopodia represented by large dorsal cirri supported by acicula. Setae include limbate setae and in some genera, bidentate hooded setae. Lysaretidae is a small family of rather large, mainly tropical shallow-water polychaetes. The general appearance of the worms is that of a lumbrinerid, but the large dorsal cirri are distinct, as is the jaw-apparatus. Imjwrtant revisions include Fauchald (1970) and Knox and Green (1972).
Key to Genera 1 a. lb. 2a (lb). 2b (lb). 3a (2b). 3b (2b). One distinct peristomial segment present Oenone Two distinct peristomial segments present 2 One short antenna present Tainokia Three antennae present 3 Proximal end of maxilla I dentate; distal end falcate Halla Proximal end of maxilla I smooth or nearly smooth; each distal end with two large fangs . . Lysarete Generic Definitions Halla Costa 1844, Nereis parthenopeia delle Chiaje 1828; 3 species. Lysaretids with two distinct peristomial rings, three antennae and the distal end of maxilla I falcate. The proximal end of maxilla I is dentate. Lysarete Kinberg 1865, L. brasiliensis Kinberg 1865; 2 species. Lysaretids with two distinct peristomial rings, three antennae and the distal end of maxilla I divided into two large fangs. Proximal end of maxilla I smooth. Oenone Savigny 1818, Aglaura fulgida Savigny 1818; 4 species. Lysaretids with one distinct peristomial ring, three antennae and usually, distally falcate maxilla I. Proximal end of maxilla I dentate. Tainokia Knox and Green 1972, T. iridescens Knox and Green 1972; only species. Lysaretids with two distinct peristomial rings, one antenna and distally falcate maxilla I. Proximal end of maxilla I is dentate. Taxonomic Notes The number of species involved in the circumtropical complex referred to as Oenone fulgida has not been determined. Crossland (1924) investigated some of the variabiHty of the jaw-apparatus, but did not have large enough materials to confirm or reject the presence of distinct sub-groups. Halla parthenopeia also has been reported from cosmotropical areas and may be another species-complex. Invalid Genera Aenone Risso n Quatrefages 1865, indeterminable Aglaura Savigny 1818, see Oenone Aglaurides Ehlers 1868, see Oenone Andromache Kinberg 1865, see Oenone Cirrobranchia Ehlers 1868, see Halla Danymene Kinberg 1865, indeterminable Larymna Kinberg 1865, indeterminable Plioceras Quatrefages 1865, see Halla
FAMILY DORVILLEIDAE CHAMBERLIN 1919C
Eunicea with two pairs of antennae. Maxillae consist of one or two series of small jaw-pieces and paired carriers; mandibles present. Notopodia reduced, but with setae and acicula in most forms. Setae include simple and composite hooks, furcate and limbate setae. Dorvilleids are mainly small polychaetes, considered most common in shallow water, but recently recovered in increasing numbers from deeper water (Jumars 1974),
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One pair of the antennae is referred to as palps; they differ in structure from the other pair, in that they often are articulated. Either one or both pairs may be reduced.
The jaw-apparatus may also be reduced in some forms. Jumars (1974) has revised the generic classification of the family.
Key to Genera 1 a. lb. 2a (la). 2b (la). 3a (lb). 3b (lb). 4a (3a). 4b (3a). 5a (3b). 5b (3b). 6a (5b). 6b (5b). 7a (6b). 7b (6b). Notacicula present Notacicula absent Furcate setae present Furcate setae absent Furcate or geniculate setae present Furcate or geniculate setae absent Palps well developed Second pair of antennae reduced or absent Only simple acicular setae present Both capillary and compound setae present Setae of first seliger markedly different from the others Setae of first seliger similar to others First antennae long and cirriform First antennae reduced and papilliform 2 3 Schistomeringos Dorvillea 4 5 Protodorvillea Meiodorvillea Parophryotrocha 6 Exallopus 7 Apophryotrocha Ophryotrocha
Generic Definitions Apophryotrocha Jumars 1974, A. mutabiliseta Jumars 1974; only species. Dorvilleids with very long, cirriform antennae and well-developed palps. Dorsal cirri without acicula. Setae of two kinds, capillaries and composite heterogomph falcigers. Both pairs of carriers and the four basal plates of the jaws fused into a single structure. Dorvillea Parfitt 1866, Staurocephalus rubrovittatus Grube 1855; 15 species. Dorvilleids with both antennae and palps well developed and of the same length. Dorsal cirri with acicula. Setae include capillaries and heterogomph falcigers. Carriers and four rows of denticles always present; carriers may be variously fused to the basal plates. Exallopus Jumars 1974, E. cropion Jumars 1974; only species. Dorvilleids with both antennae and palps well developed and of the same length. Dorsal cirri absent. Setae include capillaries and composite heterogomph falcigers and in the first seliger, large, curved composite hooks. One pair of carriers and two rows of denticles present; carriers fused to the basal plates. Meiodorvillea Jumars 1974, Protodorvillea minuta Hartman 1965; 3 species. Dorvilleids with small clvate antennae, and small palps, which may be absent. Dorsal cirri small or absent, never with acicula. Setae include capillaries, furcate setae and composite heterogomph falcigers. Carriers and two rows of denticles present.
Ophryotrocha Claparde and Mecznikow 1869, O. puerilis Claparde and Mecznikow 1869; 11 species. Dorvilleids with antennae and palps reduced to papilliform projections. Dorsal cirri without acicula, may be absent. Setae include capillaries and heterogomph falcigers. Carriers and basal plates fused into a characteristic ice-tong shape. Four rows of denticles present. Parophryotrocha Hartmann-Schrder 1971, Ophryotrocha isochaeta Eliason 1962; only species. Dorvilleids with antennae and palps absent. Dorsal cirri absent. Only simple, acicular setae present. Four rows of denticles present, carriers and basal plates fused into one piece. Protodorvillea Pettibone 1961, Staurocephalus kefersteini Mclntosh 1869; 5 species.
FIGURE 32. Family DORVlLLEmAE, Dorvillea articlala, Newport, California, shallow subtidal, 50x.
1976
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Dorvilleids with well-developed palps; antennae small or absent. Dorsal cirri without acicula. Setae include capillaries, furcate setae and composite heterogomph setae. Carriers and four rows of denticles present; carriers sometimes fused with basal plates. Schistomeringos Jumars 1974, Nereis rudolphii delle Chiaje 1828; 10 species. Dorvilleids with well-developed antennae and palps of approximately the same length. Dorsal cirri with acicula. Setae include capillaries, furcate setae and composite heterogomph falcigers. Four rows of denticles present; carriers may be fused with basal plates. Taxonomic Notes The genera of this family have been confused to a considerable extent, as noted by Pettibone (1961) and Jumars (1974), Thanks to these two surveys, the matter seems to have been adequately clarified. Invalid Genera Anisoceras Grube 1856, see Dorvillea and indeterminable
Eleonopsis Esmark 1878, see Ophryotrocha Prionognathus Keferstein 1862, see Schistomeringos Staurocephalus Grube 1855, see Dorvillea Stauroceps Verrill 1900, see Dorvillea Stauronereis Verrill 1900, see Dorvillea and Schistomeringos Telonereis Verrill 1900, see Dorvillea The following two families are considered here free-standing, unrelated families of the order Eunicida. They are both very small, in terms of numbers of species, and parasitic on decapod crustaceans and fishes, respectively. Probably in response to this habit, they have been modified so that the only characters they have in common with other members of the order, is the structure of the jaw-apparatus, which makes them resemble members of the Eunicida more closely than they resemble members of any other jawed family.
FAMILY HISTRIOBDELLIDAE VAILLANT
1890
Eunicida with five antennae; one pair of lower jaws and a single lateral jaw. One pair of anterior and one pair of posterior appendages always present; a varying
33. (A), Family ICHTHYOTOMIDAE, Ichthyotomus sanguinarias, combined from several sources, ventral view, about 30x; (B), Family HISTRIOBDELLIDAE, Histriobdella homari, from Homarus americanus, Woods Hole, Massachusetts, 95x; (C), Family STERNASPIDAE, Siernaspis scutata, off Santa Catalina Island. 23 m, anterior end inverted, 5x; (D), the above with the anterior end everted, lOx.
FIGURE
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number of lateral appendages also present. Setae absent. Parasitic on replant decapod crustaceans. Histriobdellids are known as parasites in the branchial chambers of crustaceans, both from fresh-water and
marine environments. They are very small, and that, combined with their habitat makes it likely that they are considerably more common than the few scattered records indicate.
Key to Genera la. lb. With a single pair of lateral appendages With at least two pairs of lateral appendages Histriobdella Stratiodritus Renier 1807. The total number of currently recognized species is about ten. The sternaspids are among the most easily recognized polychaetes with the usually dark yellow or reddish chitinized shield. They are common in sandy and muddy substrates in all depths, but are perhaps most usually found in about 100-200 m depth. Sternaspids are only rarely found in large numbers. They are burrowers in the sand and mud. Invalid Genera Eckinorhynchus Jfivst; Renier 1804, see. Stemaspis Schreiberius Otto 1821, see Stemaspis Thalassema Ranzani 1817, see Stemaspis ORDER OWENIIDA Prostomium fused to the anterior segments; prostomium sometimes produced in lobes or as a folded membrane; proboscis a muscular pad. Neuropodial hooks in dense fields.
FAMILY OWENIIDAE RIOJA
The genera are defined as indicated in the key. Each genus is known in just a few species. Histriobdella van Beneden 1858, H. homari van Beneden 1858, only species. Stratiodrilus Has well 1900, 5. tasmanicus Has well 1900, 4 species. Invalid Genus Histriodrilus Foettinger 1884, see. Histriobdella
FAMILY
IcHTHYOTOMtDAE
EISIG
1906
Eunicida with one antenna, one pair of lateral jaws. Notopodia with acicula, but otherwise asetigerous, neurosetae composite. Parasitic on fishes. Ichthyotomus is known for a single species, /. sanguinarius Eisig 1906 from the Gulf of Naples. It is parasitic on the fins of eels. Fauvel (1958) gives a good description and illustration of the form; the illustration given here has been redrawn from that illustration. ORDER STERNASPIDA Posterior ventrum covered by a stiff, chitinized, mineral-impregnated shield. Eversible pharynx axial, can be inverted with the first three setigers.
FAMILY STERNASPIDAE CARUS
1917
1863
Short-bodied polychaetes with indistinct segmentation; prostomium without appendages. All setae simple, those in the first three setigers thick, falcate spines; those associated with the shield, slender capillanes. The family is known for one genus, Stemaspis Otto 1821, with type species, Eckinorhynchus scutatus
Body cylindrical with long anterior segments and short posterior ones; tubicolous. Notosetae capillary, neurosetae very small bi- or tridentate hooks in dense fields. The oweniids are characteristically rather small, tubicolous animals, the tubes are usually short, and they are often capable of moving around with the tube. They have turned out to be quite frequently reported from moderate depths on the continental slopes, but do not appear to be common in abyssal depths. The shape of the rather characteristic small hooks has been well demonstrated by Thomassin and Picard (1972) with help of scanning electron microscope.
Key to Genera la. lb. 2a (la). 2b (la). 3a (lb). Prostomium Prostomium Prostomium Prostomium Prostomium anteriorly produced into a low collar or tentacular crown rounded or bilobed forming a low collar, ventrally deeply incised forming a tentacular crown, ventrally enth-e deeply bilobed with paired palps 2 3 Galathowenia Owenia Myriowenia
i976 3b (lb). 4a (3b). 4b (3b).
115 4 Myriochele Myrioglobula
Prostomium rounded, palps absent First two or three setigers with notosetae only Only first setiger with notosetae only
Generic Definitions Galathowenia Kirkegaard 1959, G. africana Kirkegaard 1959; only species. Prostomium anteriorly produced into a low collar that ventrally is deeply incised with one lobe overlapping the other. Eyes present. First three setigers without neurosetae. Myriochele Malmgren 1867, M. heeri Malmgren 1867; 12 species. Prostomium anteriorly rounded. First two or three setigers with notosetae only. Myrioglobula Hartman 1967, M. antrctica Hartman 1967; only species. Prostomium anteriorly rounded. Only one anterior setiger with notosetae only. Myriowenia Hartman 1960, M. californiensis Hartman 1960; 2 species. Prostomium deeply bilobed with a pair of grooved palps attached anteriorly. First three setigers without neurosetae. Owenia delle Chiaje 1841, O. fusiformis delle Chiaje 1841; 11 species. Prostomium produced anteriorly into a low tentacular crown with flattened lobate projections. Three first setigers without neurosetae. Invalid Genera Ammochares Grube 1847, see Owenia Mitrara Mller 1851, larval forms Ops Carrington 1865, set Owenia Psammocollus Grube 1868, sec Myriochele
ORDER FLABELLIGERIDA
Anterior pharynx with ventrolateral muscular pad or unreinforced. Anterior end retractable within a sheath formed by the first setigers.
FAMILY FLABELLIGERIDAE SAINT-JOSEPH
1894
34. (A), Family OWENIIDAE, Owenia collaris, combined from several sources. 12.5x; (B), Family FLABELLIGERIDAE, Pherusa inflata. after Hartman, 1969, 5x; (C), Family POEOBIIDAE, Foeobius meseres, redrawn and simplified after Robbins, 1965, about 5x; (D), Family FAUVELIOPSIDAE, Fauveliopsis brevis, after Hartman and Fauchald, 1971. 12x.
FIGURE
Body cylindrical or fusiform, most forms with epidermis covered by papillae. Pro- and peristomia retractable between the first three setigers. Prostomium a slender ridge with palps at the posterolateral sides. Peristomium with an expanded dorsal membrane cîrrying branchiae. Notosetae cross-barred or smooth, and slender; neurosetae either similar or more thickened
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spines or composite with falcate, unidentate or bidentate appendages. The flabelligerids characteristically are heavily impregnated with sand or mud in a matrix of mucus; in some forms this mucus forms a complete smooth encasing for the animal {Flabeiligera), but in most each
individual papilla, which secretes the mucus, has its separate cover of particles. Dissection of the retractable anterior end is necessary for safe identification of the several similar genera. The number and structure of the branchiae and the structure of the branchial membrane are important identificatory characters.
Key to Genera la. lb. 2a (lb). 2b (lb), 3a (2a). 3b (2a). 4a (2b). 4b (2b). 5a (4b). 5b (4b). 6a (5a). 6b (5a). 7a (5b). 7b (5b). 8a (7a). 8b (7a). 9a (7b). 9b (7b). 10a (9a). 10b (9a). 1 la (9b). lib (9b). I2a (11 b). 12b (lib). 13a (12b). 13b (I2b). Body with a distinct incision just posterior to the setigers carrying the cage-forming setae Therochaeta Body without distinct incisions 2 Neurosetae composite or pseudocomposite 3 Neurosetae entirely simple, but usually distinctly cross-barred 4 Body encased in a smooth continuous mucus sheath Flabeiligera Body with individual papillae covered with mucus and impregnated with debris Flabelliderma Branchiae absent Bradabyssa Branchiae present 5 Branchial membrane long, tonguelike, sometimes doubled 6 Branchial membrane short, rounded or triangular 7 Branchial membrane club-shaped, with branchial filaments attached distally on all sides .. Coppingeria Branchial membrane flattened, with branchial filaments attached on one side only Piromis All setae capillary 8 At least some neurosetae acicular or falcigerous 9 Body anteriorly inflated with tapering posterior end Diplocirrus Body, short, flattened and nearly disc-shaped llyphagus Cephalic cage poorly developed or absent 10 Cephalic cage well developed 11 Neurosetae distinctly thicker than notosetae, four pairs of branchiae Trophoniella Neurosetae only slightly thicker than notosetae; numerous pairs of branchiae Brada Body covered by a thick mucus sheath Buskiella Body not covered by a mucus sheath, often sand-incrusted 12 A long oral tube present Therochaetella Oral tube absent 13 Notosetae serrated and plumose Pantoithrix Notosetae cross-barred capillaries Pherusa Generic Definitions Brada Stimpson 1854, B. granosa Stimpson 1854; 21 species. Body short and fusiform; cephalic cage absent. Numerous branchial filaments on a short branchial membrane. Neurosetae slightly thicker than notosetae, distally acicular. Bradabyssa Hartman 1967, B. papillota Hartman 1967; only species. Body short and fusiform; cephalic cage absent. Branchiae absent; branchial membrane short and folded. Neurosetae simple hooks. Buskiella Mclntosh 1885, B. abyssorum Mclntosh 1885; 2 species. Body anteriorly inflated; mucus sheath covering the whole body. Cephalic cage present; parapodia very
prominent. Oral tube present. Branchial membrane triangular with numerous branchial filaments. Neurosetae acicular. Coppingeria Haswell 1892, C. longisetosa Haswell 1892; only species. Body anteriorly inflated; cephalic cage present. Branchial membrane long, cylindrical and slender, with the branchial filaments attached on all sides distally. Neurosetae simple hooks. Diplocirrus Haase 1915, Trophonia glauca Malmgren 1867; 7 species. Body anteriorly inflated; cephalic cage present in some forms. Four pairs of branchiae of two kinds on a short branchial membrane. All setae capillaries. Flabelliderma Hartman 1969, Flabeiligera commensalis Moore 1909; only species.
1976
m
Invalid Genera
Body cylindrical; body papillae covered with thick mucus and encrusted with debris. Cephalic cage present. Pseudocomposite hooks present in most neuropodia. Flabelligera Sars 1829, F. affinis Sars 1829; 18 species. Body cylindrical; covered completely by smooth mucus sheath. Cephalic cage present; pseudocomposite or composite hooks in the neuropodia. Ilyphagus Chamberiin 1919c, /. bythincola Chamberlin 1919c; 11 species. Body stout, flattened and nearly disc-shaped, covered with large papillae and mud- or sand-incrusted in most forms. Cephalic cage present in some forms. Four pairs of branchiae on a short branchial membrane. All setae capillaries. Pantoithrix Chamberlin 1919c, Pherusa chilensis Schmarda 1861; only species. Body anteriorly inflated; cephalic cage present. Notosetae serrated and plumose capillaries; neurosetae bidentate hooks. Six pairs of branchiae. Pherusa Oken 1807, Amphitrite plumosa O,F. Mller 1776; 43 species. Body anteriorly inflated; cephalic cage present. Either four or many pairs of branchiae on a short branchial membrane. Most neurosetae uni- or bidentate hooks. Piromis Kinberg 1867b, P. arenosus Kinberg 1867b; 10 species. Body anteriorly inflated; cephalic cage present. Branchial membrane prolonged, flattened and tongueshaped, either single or double, with numerous branchial filaments. Neurosetae mostly uni- or bidentate hooks. Therochaeta Chamberlin 1919c, Stylarioides coUarifer Ehlers 1887; 6 species. Anterior end inflated; cephalic cage present. A distinct incision present behind the last setiger that carries the cage setae. First post-incisional setiger with series of enlarged, usually anteriorly directed papillae. Composite hooks present on some anterior setigers in most species; otherwise simple neuropodial hooks. Therochaetella Hartman 1967, T. chilensis Hartman 1967; only species. Anterior end inflated; cephalic cage present. Long narrow oral tube present. Numerous pairs of branchiae on a short branchial membrane. Neurosetae distally bifid and falcate. Trophoniella Caullery 1944, T. avicularia Cauilery 1944; 3 species. Anterior end inflated; cephalic cage very poorly developed or absent. Four pairs of branchiae on a short branchial membrane. Neurosetae bi- or unidentate hooks.
Aristenia Savigny in Quatrefages 1865, indeterminable Balanochaeta Chamberlin 1919c, see Pherusa Chloraema Dujardin 1839a, see Flabelligera Flemingia Johnston 1846, sec Pherusa Lophocephalus Costa 1841, see Pherusa Pycnoderma Grube 1877, ses Piromis Saphobranchia Chamberlin 1919c, see Diplocirrus Semiodera Chamberlin 1919c, see Piromis Siphonostoma Rathke 1843, se.e Flabelligera Siphostoma Otto 1821, see Flabelligera Stylarioides delle Chiaje 1841, see Pherusa Tecturella Stimpson 1854, see Flabelligera Trophonia Cuvier 1830, see Pherusa Zorus Webster and Benedict 1887, indeterminable
FAMILY POEOBIIDAE HEATH
1930
Body saclike without external segmentation or setae. Anterior end fused to the rest of the body; containing a circlet of eversible tentacles. Two distinct septa only polychaete characters. Pelagic. The poeobiids are considered related to the flabelligerids because the anterior end is retractable, and because they have chlorocruorin as one of their pigments (Robbins 1965). The family contains a single genus and species, Poeobius meseres Heath 1930. Incertae Sedis Enigma Betrem 1924, E. terwillei Betrem 1924; only species. Parapodia and setae absent, retractile branchiae and palps present; one septum observed, pelagic. This form resembles the poeobiids in several respects, it has been reported only once, but unfortunately the original material has been lost (Hartman 1967). ORDER FAUVELIOPSIDA Pro- and peristomium without appendages. Pharynx with a ventral muscular pad. All setigers biramous with simple limbate setae and a small rounded papilla between the rami. These small, deep-water polychaetes are incompletely known. They were grouped formerly with the flabelligerids from which they differ in that they lack the papillar investments, the retractable anterior end and the characteristic setae. The shape of the prostomium also is markedly different. Further information may demonstrate that they are related to other groups of polychaetes; a separate order appears justified for the time being.
FAMILY FAUVELIOPSIDAE HARTMAN
1971
Small, smooth-bodied polychaetes without anterior appendages. Proboscis a ventral muscular pad. All
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setigers biramous with simple limbate setae and a small rounded papilla between the rami. Parapodial lobes reduced. The family was erected by Hartman (1971) for Fauveliopsis Mclntosh, with type F. challengeriae Mclntosh 1922, as well as for Flabeliigella Hartman 1965, Flota Hartman 1967 and Bruunilla Hartman 1971. Flabeliigella was shown to belong to the Acrocirridae by Orensanz (1974b) and are cited under that family above. Flota and Bruunilla differ sharply from Fauveliopsis and are characterized here as free-standing genera without obvious familial affiliations. The whole family is then reduced to the type-genus, Fauveliopsis, with eight described species. Incertae Sedis Bruunilla Hartman 1971, B. natalensis Hartman 1971; only species. Prostomium small and triangular attached ventral to the large peristomium. Peristomium with five tentacular cirri. Paired short palps lateral to the mouth as flat pads. Prominent lateral parapodia present; all biramous, both rami distally strongly pointed, with embedded acicula, setae absent. Dorsal and ventral cirri present on most segments. Flota Hartman 1967, F. flabelligera Hartman 1967; only species. Body with less than ten segments; short and flattened. Prostomium a simple conical lobe with a pair of tri-
lobed processes. Pharynx muscular and eversibje, scoop-shaped and open dorsally. Parapodia biramous with setae of two kinds; slender smooth acicular rods and thicker and cross-barred acicular setae. Body with papillae, especially on the parapodia. Encased in a thick mucus sheath, pelagic. ORDER TEREBELLDDA Prostomium without appendages; peristomium with a series of feeding appendages; pharynx with a ventral muscular pad. At least one pair of branchiae present (rarely absent).
FAMILY SABELLARIIDAE JOHNSTON
1865
Tubicolous polychaetes with the body in three regions; posterior region an asetigerous anal tube, ftostomium a narrow ridge fused laterally to the first setiger. Setae of first setiger forming an operculum with setae in one to three rows. Thorax with two rudimentary segments and three or four parathoracic setigers. Median region with notopodial pectinate uncini and ventral capillary setae. The sabellariids are important as reef-builders in certain areas (Schfer 1972). All of them are tubebuilders and tend to build on firm substrates; consequently, most of them are present in shallow water only, but members of two genera (Phalacrostemma and Monorchos) appear to be most common in slope depths.
Key to Genera la. I b. 2a (la). 2b (la). 3a (lb). 3b (lb). 4a (3a). 4b (3a). 5a (4b). 5b (4b). 6a (3b). 6b (3b). With a single row of paleae in the operculum 2 With at least two rows of paleae in the operculum 3 The prolonged opercular peduncles free from one another Phalacrostemma Opercular peduncles short and fused Monorchos Two rows of paleae 4 Three rows of paleae 6 Hooks absent Gunnarea A pair of large hooks dorsal and proximal to the opercular paleae 5 Three parathoracic setigers Idanthyrsus Four parathoracic setigers Lygdamis Middle opercular paleae cover the inner ones; operculimi flattened cone Phragmatopoma Middle opercular paleae not concealing the inner ones; operculum open and bristly Sabellaria Generic Definitions Gunnarea Johansson 1927, Hermella capensis Schmarda 1861; only species. Two rows of paleae; opercular peduncles fused; hooks and accessory setae absent on operculum. Three parathoracic setigers. Idanthyrsus Kinberg 1867b, /. armatus Kinberg 1867b; 5 species, Two rows of paleae; opercular peduncles fused; hooks present dorsal and proximal to the paleae; accessory setae absent. Three parathoracic setigers. Lygdamis Kinberg 1867 b, L. indicus Kinberg 1867b; 12 species. Two rows of paleae; opercular peduncles long and separated; hooks present dorsal and proximal to the paleae; accessory setae absent. Four parathoracic setigers.
1976
119
c^\
?n
OA
FIGURE
35. (A), Family PECTINARllDAE, Amphictene capensis, after Day, 1967, 3x; (B), pectinariid-tube, 2x; (C), Family SABELLARIIDAE, tube pattern of Phragmatopoma sp. natural size; (D), Phragmatopoma califomica, after Hartman, 1969, 5x.
Monorchos Tread well 1926, M. philippinensis Treadwell 1926; 2 species, A single row of paleae; opercular peduncles fused; hooks present dorsal and proximal to the paleae and two rows of accessory setae between the paleal rows. Three parathoracic setigers. Phalacrostemma Marenzeller 1895, P. cidariophilwn Marenzeller 1895; 5 species. A single row of paleae; opercular peduncles long and separated; hooks present dorsal and proximal to the
paleae; accessory setae absent. Four parathoracic setigers. Phragmatopoma Mrch 1863, P. caudate Morch 1863; 8 species. Three rows of paleae; opercular peduncles short and fused; hooks and accessory setae absent. Three parathoracic setigers. Middle opercular paleae covers the inner ones; operculum conical. Sabellaria Savigny 1818, Sabella alveolata Linnaeus 1767; 28 species.
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Three rows of paleae; opercular peduncles short and fused; hooks absent, accessory setae sometimes present. Three parathoracic setigers. Middle opercular paleae pointed distad, operculum open and generally rather bristly in appearance. Taxonomic Notes The "accessory setae" referred to in the above definitions, include setae associated with the operculum; they usually are acicular in appearance and may be present between the rows of paleae (Monorchos) or proximal to the paleae proper (Sabellaria). The family is under current revision by Dr. David Kirtley and the numbers of species assigned to each genus may be changed drastically. The generic sub-division appears reasonably stable. Invalid Genera Centrocorone Grube 1850, see Sabellaria Chrysodon Okenin Quatrefages 1865, see Sabellaria Cryptopomatus Gravier 1908, see Idanthyrsus
Eupallasia Augener 1927a, see Lygdamis Hermella Savigny 1818, see Sabellaria Pallasia Quatrefages 1848, see Idantkyrsus Pallasina Annenkova 1925, see daithyrsus Tetreres Caullery 1913, see Lygdamis
FAMILY PECTINARIIDAE QUATREFAGES
1865
Body separated into three regions, including thorax, abdomen and a posterior scaphe. Prostomium reduced. First seliger with expanded, strong setae (paleae) forming a comb used in digging. Other setae include short capillaries, pectiniform uncini and scaphal spines. Tubes short, more or less tusk-shaped, built of relatively large particles. The pectinariids are among the most characteristic polychaetes with their strongly golden, coppery or brassy paleal setae and the gently curved, tapering, tusk-shaped, but usually very fragile tubes. The family often is recognized for only two genera, Petta and Pectinaria, in which case the other listed genera are considered subgenera of Pectinaria.
Key to Genera la. lb. 2a (lb). 2b (lb). 3a (2b). 3b (2b). 4a (3b). 4b (3b). Cephalic veil marginally smooth, scaphe not distinctly separated fiwm abdomen Cephalic veil marginally cirrate; scaphe distinctly set off from abdomen Twelve uncinigers, cephalic veil at least partly fused to the operculum Thirteen uncinigers, cephalic veil completely free from the operculum Opercular rim cirrate Opercular rim smooth Uncini with major teeth in a single row Uncini with major teeth in two rows Petta 2 Lagis 3 Amphictene 4 Cistenides Pectinaria
Generic Definitions Amphictene Savigny 1818, Amphitrite auricoma O.P. Mller 1776; 8 species. Cephalic veil free from operculum, marginally cirrate; opercular rim marginally cirrate. Scaphe distinctly set off from abdomen. Uncini with major teeth in double rows; 13 uncinigers. Cistenides Malmgren 1866, Sabella granlala Linnaeus 1767; 8 species. Cephalic veil free from operculum, marginally cirrate; opercular rim marginally smooth. Scaphe distinctly set off from abdomen. Uncini with major teeth in a single row, 13 uncinigers present. Lagis Malmgren 1866, L. koreni Malmgren 1866; 8 species. Cephalic veil at least partly fused to operculum, marginally cirrate; opercular rim marginally smooth. Scaphe
distinctly set off from abdomen. Uncini with major teeth in two or more rows; 12 uncinigers present. Pectinaria Savigny 1818, Nereis cylindraria blgica Pallas 1766; 18 species. Cephalic veil free from operculum, marginally cirrate. Opercular rim marginally smooth. Scaphe distinctly set off from the abdomen. Uncini with major teeth in double rows; 13 uncinigers present. Petta Malmgren 1866, P. pusilla Malmgren 1866; 4 species. Cephalic veil free from operculum; marginally smooth. Opercular rim marginally cirrate. Scaphe not distinctly set off firom abdomen. Uncini with major teeth in a single row; 14 uncinigers. Invalid Genera Ariapithes Kinberg 1867b, indeterminable Cistena Leach 1816, see Pectinaria
i976 Labiaria Sveshnikov 1959, larval forms Scalis Grube 1846, indeterminable
111
FAMILY AMPHARETIDAE MALMGREN
1867
Body with two regions; anterior region with biramous parapodia; posterior region with well-developed neuropodia, notopodia reduced or absent. Prostomium simple or complex with lateral folds and glandular ridges. Two to four pairs of smooth or pinnate branchiae present. Notopodia! capillary setae present in thorax, neuropodial uncini present both in thorax and abdomen. Thoracic uncini with major teeth in one or a few rows, rarely crested; abdominal uncini similar, but more frequently crested. Nuchal hooks and anterior acicular setae present in some forms. The ampharetids resemble the terebellids; the main feature used to separate the two families is behavioral in that the former will withdraw the buccal tentacles completely within their mouth; the latter do not. Addition-
ally, the ampharetids usually have a few pairs of simple 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 and nearly always distinctly crested in the terebellids. The ampharetids have turned out to be very common in deep water and a whole mass of previously unrecognized genera have been reported from deep water over the last ten years. The major classification was reviewed by Day (1964) who reduced greatly the number of monotypic genera. The present review retains most of the genera Day fused; it is not clear that the characters Day used to identify his genera are any more precise than those he disregarded. Since a large number of additional taxa are now being described, it appears best to await further fusions of genera, until the current deep-water material has been worked up in detail. The genus Oeorpata is not clearly separable from Isolda according to Day (1964) and is incompletely known. It has been included in the definitions, but not in the key.
Key to Genera la. lb. 2a ( I a). 2b (la). 3a (2a). 3b (2a). 4a (3a). 4b (3a). 5a (3b). 5b (3b). 6a (2b). 6b (2b). 7a (6a). 7b (6a). 8a (6b). 8b (6b). 9a (8b), 9b (8b). 10a (lb). lOb (lb). 1 la (lOa). lib (lOa). 12a (lOb). 12b (10b). 13a (12b). 13b (12b). 14a (13b). MELINNINAE 2 First severed neuropodia without acicular setae; all neurosetae uncini AMPHARETINAE 10 Nuchal hooks present 3 Nuchal hooks absent 6 Branchiae smooth 4 Branchiae in part papillose or pennate 5 Two pairs of nuchal hooks; 12 thoracic uncinigers Moyanas A single pair of nuchal hooks, 14 thoracic uncinigers Melinna Four pairs of branchiae, two smooth and two papillose; capillary notosetae on setigers 3-4 Isolda Three pairs of branchiae, one smooth, two papillose; notosetae absent on setigers 3-4 Irana Ten thoracic uncinigers present 7 At least 12 thoracic uncinigers present 8 Three pairs of branchiae Melinnopsides Four pairs of branchiae Melinnopsis Sixteen thoracic uncinigers present; all buccal tentacles similar Melinantipoda Maximally 14 thoracic uncinigers present; one or two buccal tentacles very large 9 Segment 6 with a distinct dorsal crest between the notopodia Melinnexis A dorsal crest absent Amelinna At least three pairs of branchiae 11 Two pairs of branchiae 12 Three pairs of branchiae present 15 Four pairs of branchiae 35 Fourteen thoracic uncinigers; paleae present Ecamphicteis Maximally 12 thoracic uncinigers; paleae absent 13 Nine uncinigers, first uncini in a short row similar to all others Egamella Twelve uncinigers, first uncini in a long row 14 Prostomium anterioriy pointed Auchenoplax First several neuropodia with fine acicular setae; other neuropodia with uncini
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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 straemi, combined from several sources, about 5x; (D), Family TEREBELLIDAE, Neoamphitrile, oeai johrtstoni, modified from live sketch, 3x.
FIGURE
1976 14b (13b). 15a (11a). 15b (1 la). 16a (15a). 16b (15a). 17a (16b). 17b (16b). 18a (15b). 18b (15b). 19a (18a). 19b ( 18a). 20a (19b). 20b (19b). 21a (20b). 21b (20b). 22a (21b). 22b (21b). 23a (22a). 23b (22a). 24a (23b). 24b (23b). 25a (18b). 25b (18b). 26a (25b). 26b (25b). 27a (26b). 27b (26b). 28a (27a). 28b (27a). 29a (28b). 29b (28b). 30a (29b). 30b (29b). 31a (27b). 31b (27b). 32a (31a). 32b (31a). 33a (31b). 33b (31b). 34a (33b). 34b (33b), 35a (1 lb). 35b (lib). 36a (35a). 36b (35a). 37a (36b). 37b (36b). 38a (37a). 38b (37a). 39a (37b). 39b (37b). 40a (39a). 40b (39a). 41a (40a). 41b (40a).
123
Prostomium anteriorly truncate Melinnoides All branchial pairs arranged in a distinctly segmental fashion 16 Only last pair of branchiae clearly associated with a specific segment 18 Eleven thoracic uncinigers; tentacular membrane large Ampharana At least 14 thoracic uncinigers; tentacular membrane small 17 First setae on segment 2; 14 thoracic uncinigers Neopaiwa First setae in segment 3; 15 thoracic uncinigers Weddellia Paleae present 19 Paleae absent 25 Nine thoracic uncinigers; last thoracic notopodia elevated Mugga Ten or more thoracic uncinigers 20 Flange present between last thoracal notopodia; ten thoracic uncinigers Melinnaia Flange or crest, if present, anterior in position; at least 11 thoracic uncinigers 21 Few stout buccal tentacles present; 11 thoracic uncinigers Amythasides Buccal tentacles, if present, numerous, 12 thoracic uncinigers 22 Dorsal crest present on one anterior setiger 23 Dorsal crest absent Eclysippe Oral membrane broadly folded, 15 thoracic setigers Eusamythella Oral membrane short; 14 thoracic setigers 24 Dorsal crest on segment 4 . Neosamytha Dorsal crest on either segment 6 or 7 Melirmampharete Ten thoracic uncinigers Muggoides At least 11 thoracic uncinigers 26 Eleven thoracic uncinigers Glyphanostomum At least 12 thoracic uncinigers 27 Twelve thoracic uncinigers 28 At least 13 thoracic uncinigers 31 At least one pair of branchiae basally flanged Samythella All branchiae cylindrical 29 Buccal tentacles papillose Neosabellides Buccal tentacles smooth 30 Oral membrane greatly prolonged Pabits Oral membrane short Eusamytha Thirteen thoracic uncinigers 32 Fourteen thoracic uncinigers 33 Notopodia of setiger 13 elevated with hirsute notosetae Sosanella Notopodia of setiger 13 at the same level as all others; hirsute setae absent Alkmaria Oral membrane smooth and folded Amythas Oral membrane tentaculate 34 Glandular ridges on prostomium Samythopsis Glandular ridges absent Samytha Paleae present (i.e. setae present on segment 3, as the first notosetae in the body) 36 Paleae absent (i.e. setae absent on segment 3 or setae already present from segment 2) 52 Flanged branchiae present on four successive segments Phyllampharete Branchiae flanged, lamellate, pennate or cylindrical; only one pair clearly associated with a segment 37 Eleven thoracic uncinigers 38 At least 12 thoracic uncinigers 39 Branchiae pennate, buccal tentacles papillose Pterampharete Both branchiae and buccal tentacles smooth Sabellides Twelve thoracic uncinigers present 40 At least 13 thoracic uncinigers present 44 Buccal tentacles papillose 41 Buccal tentacles smooth 42 Setiger 11 with notopodia elevated and notosetae distally hirsute Anobothrella Setiger 11 with notopodia at the same level as all other setigers; notosetae limbate or capillary Ampharete
124 Ala. (40b). 42b (40b). 43a (42b). 43b (42b). 44a (39b). 44b (39b). 45a (44a). 45b (44a). 46a (45b). 46b (45b). 47a (44b). 47b (44b). 48a (47a). 48b (47a). 49a (47b). 49b (47b). 50a (49a). 50b (49a). 51a (49b). 51b (49b). 52a (35b). 52b (35b). 53a (52b). 53b (52b). 54a (53a). 54b (53a). 55a 55b 56a 56b 57a 57b 58a 58b 59a 59b 60a 60b (53b). (53b). (55a). (55a). (56b). (56b). (55b). (55b). (58a). (58a). (58b). (58b).
Science Series 28
Seliger 14 with elevated notopodia and hirsute notosetae Sosane Setiger 10 or 11 with elevated notopodia and tapering or hirsute notosetae 43 Notopodial rudiments present in abdomen Anobothrus Notopodia! rudiments absent from abdomen Sosanides Thirteen thoracic uncinigers ; 45 Fourteen thoracic uncinigers 47 One pair of branchiae pennate, others smooth Pterolysippe All branchiae smooth 46 Glandular ridge on prostomium Hypania Glandular ridge absent Lysippe Two of the four pairs of branchiae lamellate 48 All four pairs of branchiae cylindrical 49 Paleae large, abdominal notopodial rudiments absent Pkyllamphicieis Paleae small, abdominal notopodial rudiments present Lysippides Glandular ridges on prostomium 50 Glandular ridges absent 51 Abdominal notopodial rudiments present Amphicieis Abdominal notopodial rudiments absent Parhypania Abdominal notopodial rudiments present Paiwa Abdominal notopodial rudiments absent Hypaniola Branchiae arranged in oblique series directly associated with distinct segments Mexamage Only the last pair of branchiae clearly associated with a segment 53 At least three pairs of branchiae lamellate 54 All branchiae cylindrical 55 All four pairs of branchiae lamellate; anus surrounded by a circle of papillae Phyllocomus Three pairs of lamellate and one pair of cylindrical branchiae; anus with two pairs of anal cirri .... Schistocomus Eleven thoracic uncinigers 56 At least 12 thoracic uncinigers 58 First two notopodia (segments 4 and 5) asetigerous , Paramage All anterior notopodia with setae 57 One pair of anal cirri Grubianella Two pairs of anal cirri Amage Twelve thoracic uncinigers 59 Fourteen thoracic uncinigers 60 Buccal tentacles papillose; all notosetae capillary Asabellides Buccal tentacles smooth; last thoracic notosetae modified Sosanopsis Buccal tentacles papillose; notopodial cirri present Paramphicteis Buccal tentacles smooth; notopodial cirri absent Amphisamytha Generic Definitions MELINNINAE. Four pairs of smooth branchiae; notosetae absent in segments 3 and 4. Nuchal hooks absent; one or two large oral tentacles present in addition to numerous smaller ones. Twelve or 13 thoracic uncinigers. Ampharana Hartman 1967, A. antrctica Hartman 1967; only species. AMPHARETINAE. Three pairs of smooth branchiae on three successive segments. Oral membrane large and folded with buccal tentacles in two lateral groups. Eleven thoracic uncinigers. Paleae absent, but small capillary setae present in segment 3. Ampharete Malmgren 1866, Amphicteis acutifrons Grube 1860; 27 species.
Alkmaria Horst 1919b, A. rominji Horst 1919b; only species. AMPHARETE>AE. Three pairs of smooth branchiae; no paleae. Glandular ridges absent; 13 thoracic uncinigers; no notopodial rudiments in abdomen. Amage Malmgren 1866, A. auricula Malmgren 1866; 14 species. AMPHARETENAE. Four pairs of smooth branchiae; no paleae. Glandular ridges present; 11 thoracic uncinigers; abdominal notopodial rudiments present. Two pairs of anal cirri. Amelinna Hartman 1969, A. abyssalis Hartman 1969; 2 species.
1976
THE POLVCHAETE WORMS
125
AMPHARETINAE. Four pairs of smooth branchiae; paleae present. Buccal tentacles papillose. Glandular ridges absent. Twelve thoracic uncinigers; abdominal notopodial rudiments absent. Notosetae absent in segment 4. Amphicteis Grube 1850, Amphitrite gunneri Sars 1835; 30 species. AMPHARETEîAE. Four pairs of branchiae, usually cylindrical, rarely folise; paleae present. Glandular ridges present. Fourteen thoracic uncinigers; abdoininal notopodial rudiments present. Amphisamytha Kessle 1917, A. japnica Kessle 1917; 2 species. AMPKARETINAE. Four pairs of smooth branchiae; paleae absent. Glandular ridges absent. Fourteen thoracic uncinigers. Abdominal notopodial rudiments present. Amythas Benham 1912, A. membranifera Benham 1921; only species. AMPHARETINAE. Three pairs of smooth or grooved branchiae; oral membrane folded and smooth. Paleae absent; 14 thoracic uncinigers. Abdominal notopodial rudiments absent. Amythasides Eliason 1955, A. macroglossus Eliason 1955; only species. AMPHARETINAE. Three pairs of branchiae; paleae present. Buccal tentacles few and large. Eleven thoracic uncinigers. Glandular ridges absent. Abdominal notopodial rudiments absent. Anobothrella Hartman 1967, Anobothrus antrctica Monro 1939a; only species. AMPKARETINAE. Four pairs of papillose branchiae. Buccal tentacles papillose; paleae present. Twelve thoracic uncinigers. Seliger 11 with notopodia elevated and hirsute notosetae. Anobothrus Levinsen 1884, Ampharete gracias Malmgren 1866; 8 species. AMPHARETINAE. Four pairs of smooth branchiae, paleae present. Twelve uncinigers. Glandular ridges absent; abdominal notopodia] rudiments present. Setiger 10 or 11 with notopodia elevated and modified notosetae. Asabellides Annenkova 1929, Sabellides sibirica Wirn 1883; 3 species. AMPHARETINAE. Four pairs of smooth branchiae; paleae absent. Twelve thoracic uncinigers; notosetae absent in segment 4. Buccal tentacles papillose; glandular ridges absent. Abdominal notopodial rudiments present. Auchenoplax Ehlers 1887, A. crinita Ehlers 1887; only species. AMPHARETINAE. Two pairs of smooth branchiae. Prostomium anterioriy sharply pointed. Notosetae ab-
sent in segments 3 and 4. Twelve thoracic uncinigers; first row of uncini very long and ventrally located. Ecamphicteis Fauchald 1972, E. elongata Fauchald 1972; only species. AMPHARETINAE. Two pairs of smooth branchiae on first two segments. Paleae present; 14 uncinigers. Glandular ridges and abdominal notopodial rudiments absent. Eclysippe Eliason 1955, Lysippe vanelli Fauvel 1936; only species. AMPKARETINAE. Three pairs of smooth branchiae (Day 1964: four pairs of branchiae in type species); paleae present; 12 thoracic uncinigers. Glandular ridges absent; abdominal notopodial rudiments absent. Egamella Fauchald 1972, E. quadribranchiata Fauchald 1972; only species. AMPKARETINAE. Two pairs of branchiae on two successive segments; branchial membrane high and laterally free from branchial bases. Segment 3 with small capillary setae; nine uncinigers present. Clvate notopodial abdominal rudiments present; glandular ridges absent. Eusamytha Mclntosh 1885, E. pacifica Mclntosh 1885; only species. AMPKARETINAE. Three pairs of smooth branchiae; paleae absent. Glandular ridges and abdominal notopodial rudiments absent. Twelve thoracic uncinigers. Eusamythella Hartman 1971, Eusamytha sexdentata Hartman 1967; only species. AMPHARETINAE. Three pairs of smooth branchiae; paleae present. Oral membrane broad and folded; dorsal crest across segment 5. Twelve thoracic uncinigers. Glandular ridges absent. Glyphanostomum Levinsen 1884, Samytha pallescens Thel 1879; 2 species. AMPHARETINAE. Three pairs of smooth branchiae; paleae absent. Glandular ridges absent. Eleven thoracic uncinigers. Grubianella Mclntosh 1885, G. antrctica Mclntosh 1885; only species. AMPHARETINAE. Four pairs of smooth branchiae; paleae absent. Glandular ridges present; abdominal notopodial rudiments present; one pair of anal cirri. Eleven thoracic uncinigers. Hypania Ostroumouw 1897, Amphicteis invalida Grube I860; 2 species. AMPHARETINAE. Four pairs of smooth branchiae, paleae present. Glandular ridges and abdominal notopodial rudiments present. Thirteen thoracic uncinigers. Hypaniola Annenkova 1927, Amphicteis (lAryandes) kowalewskii Grimm 1877; only species.
26
Science Series 28
AMPHARETINAE. Four pairs of smooth branchiae; small paleae present. Glandular ridges absent; 14 uncinigers. Irana Wesenberg-Lund 1949, /. heterobranchiata Wesenberg-Lund 1949; only species. MELINNINAE. Three pairs of branchiae, one smooth, other two pennate. Nuchal hooks on segment 4; dorsal crest on segment 6. Capillary notosetae first present in segment 7; 12 thoracic uncinigers. Isolda Mller 1858, /. pulchella Mller 1858; 4 species. MELINNINAE. Four pairs of branchiae; two smooth and two pennate. Nuchal hooks present; dorsal crest on segment 6. Capillary notosetae present in segments 5 and 6. Twelve or 13 thoracic uncinigers. Lysippe Malmgren 1866, L. labiata Malmgren 1866; 5 species. AMPHARETINAE. Four pairs of smooth branchiae; small paleae present. Glandular ridges absent. Abdominal notopodial rudiments present. Thirteen thoracic uncinigers. Lysippides Hessle 1917, Amphicteis fragilis Wollebaek 1912; only species. AMPHARETINAE. Four pairs of branchiae; two cylindrical, two flanged; small paleae present. Glandular ridges absent. Abdominal notopodial rudiments present. Fourteen thoracic uncinigers. Melinantipoda Hartman 1967, M, antrctica Hartman 1967; only species. MELINNINAE. Four pairs of smooth branchiae; nuchal hooks absent. First notosetae in segment 5. Dorsal crest on segment 6; 16 thoracic uncinigers. All buccal tentacles similar. Melinna Malmgren 1866, Sabellides cristata Sars 1851; 26 species. MELINNINAE. Four pairs of branchiae, nearly always smooth. Nuchal hooks present; dorsal crest on segment 6 present. Fourteen thoracic uncinigers. All buccal tentacles similar. Melinnampharete Annenkova 1937, M. eoa Annenkova 1937; 2 species. AMPHARETINAE. Three pairs of smooth branchiae; paleae present. Dorsal ridge either on segment 6 or 7. Glandular ridges absent; abdominal notopodial rudiments absent. Twelve thoracic uncinigers. Melinnata Hartman 1965, M. americana Hartman 1965; only species. AMPHARETINAE. Three pairs of branchiae; paleae present. Glandular ridges absent; ridge across dorsum on segment 4. Ten thoracic uncinigers; flange across dorsum between last thoracic notopodia.
Melinnexis Annenkova 1931, M. rctica Annenkova 1931; 8 species. MELINNINAE. Four pairs of smooth branchiae. Nuchal hooks absent; first notosetae in segment 5. Dorsal crest on segment 6. Thirteen or 14 thoracic uncinigers. One very large and numerous small buccal tentacles present. Melinnoides Benham 1927, M. nelsoni Benham 1927; only species. AMPHARETINAE. Two pairs of smooth branchiae; paleae absent; first notosetae in segment 5. Prostomium a small quadrangular lobe. Twelve thoracic uncinigers; first row of uncini long and ventrally displaced. Melinnopsides Day 1964, Melinnopsis capensis Day 1955; only species. MELINNINAE. Three pairs of smooth branchiae; nuchal hooks and dorsal crest absent. Ten thoracic uncinigers. First notosetae in segment 5; segment 6 without neurosetae. Melinnopsis Mclntosh 1885, M. atlntica Mclntosh 1885; only species. MELINNINAE. Four pairs of smooth branchiae; nuchal hooks absent; first notosetae on segment 5. Dorsal crest absent. Ten thoracal uncinigers. Buccal tentacles all similar. Mexamage Fauchald 1972, M. corrugata Fauchald 1972; only species. AMPHARETINAE. Four pairs of branchiae on four successive segments. Paleae absent; notopodia on segments 2 and 3, but not setae. Ten thoracic uncinigers; no fused anterior segments. Moyanus Chamberlin 1919c, M. explorons Chamberlin 1919c; only species. MELINNINAE. Four pairs of smooth branchiae. Nuchal hooks on both segments 4 and 5. Dorsal crest on segment 6. Twelve thoracic uncinigers. Prostomium prolonged with oral lobe suspended below it. Mugga Eliason 1955, M. wahrbergi Eliason 1955; only species. AMPHARETINAE. Three pairs of smooth branchiae; paleae present. Glandular ridges absent; abdominal notopodial rudiments absent. Nine thoracic uncinigers. Last thoracic notopodia elevated with modified notosetae. Muggoides Hartman 1965, Ai. cinctus Hartman 1965; only species. AMPHARETINAE. Three pairs of smooth branchiae; paleae absent. Glandular ridges and abdominal notopodial rudiments absent. Ten thoracic uncinigers; last thoracic notopodia elevated with modified notosetae. Neopaiwa Hartman and Fauchald 1971, N. cirrata Hartman and Fauchald 1971; only species.
1976
AMPHARETINAE. Three pairs of branchiae on three successive segments starting on segment 2. Capillary notosetae present from segment 2; paleal development absent. Fourteen thoracic uncinigers present. Glandular ridges absent; abdominal notopodial rudiments well developed. Neosabellides Hesslc 1917, N. elongata Kessle 1917; 2 species. AMPHARETINAE. Three pairs of smooth branchiae; paleae absent; first notosetae present on segment 5. Glandular ridges absent; abdominal notopodial rudiments present. Twelve thoracic uncinigers. Buccal tentacles papillose. Neosamytha Hartman 1967, N. gracilis Hartman 1967; only species. AMPHARETINAE. Three pairs of smooth branchiae; paleae present. Glandular ridges and abdominal notopodial rudiments absent. Twelve thoracic uncinigers. Dorsal ridge across segment 4. Oeorpata Kinberg 1867b, O. armata Kinberg 1867; only species. MELINNINAE. Incompletely known, not clearly separable from Isolda (cf. Day 1964). Pabits Chamberlin 1919c, P. deroderus Chamberlin 1919c; only species, AMPHARETINAE. Three pairs of smooth branchiae; paleae absent. Glandular ridges absent; notopodial rudiments present in abdomen. Twelve thoracic uncinigers. Oral lobe prolonged. Paiwa Chamberlin 1919c, P. abyssi Chamberlin 1919c; only species. AMPHARETINAE. Four pairs of smooth branchiae; small paleae present. Glandular ridges absent; small abdominal notopodial rudiments present. Fourteen thoracic uncinigers. Paramage Caullery 1944, P. madurensis Caullery 1944; only species. AMPHARETINAE. Four pairs of smooth branchiae. Paleae absent, first notosetae on segment 6; notopodial lobes present on segments 4 and 5. Glandular ridges and abdominal notopodial rudiments absent. Eleven thoracic uncinigers. Paramphicteis Caullery 1944, Sabellides angustifolia Grube 1878; only species. AMPHARETINAE. Four pairs of smooth branchiae; paleae absent. Glandular ridges present; small abdominal notopodial rudiments present. Fourteen thoracic uncinigers. Parhypania Annenkova 1928, Amphicteis brevispinus Grube 1860; only species.
AMPHARETINAE, Four pairs of smooth branchiae; paleae present. Glandular ridges present; abdominal notopodial rudiments very small. Fourteen thoracic uncinigers. Phyllampharete Hartman and Fauchald 1971, P. longicirrata Hartman and Fauchald 1971; only species. AMPHARETINAE. Four pairs of flanged branchiae on four successive segments starting on segment 2. Paleae absent; notosetae present from segment 2, Glandular ridges absent; abdominal notopodial rudiments present. Eleven thoracic uncinigers. Prostomium anteriorly incised. Phyllamphicteis Augener 1918, P. collaribranchis Augener 1918; 2 sjiecies. AMPHARETINAE. Four pairs of branchiae, two smooth and two lamellate. Paleae present. Glandular ridges and abdominal notopodial rudiments absent. Fourteen thoracic uncinigers. Phyllocomus Grube 1877, P. crocea Grube 1877; only species. AMPHARETINAE. Four pairs of flanged branchiae; paleae absent. Glandular ridges absent; abdominal notopodial rudiments present. Twelve thoracic uncinigers. Anus surrounded by a circle of papillae. Pterampharete Augener 1918, P. luderitzi Augener 1918; only species. AMPHARETINAE. Four pairs of pennate branchiae; paleae present. Segment 4 without notosetae. Glandular ridges and abdominal notopodial rudiments absent. Eleven thoracic uncinigers. Buccal tentacles papillose. Pterolysippe Augener 1918, P. bipennata Augener 1918; only species. AMPHARETINAE. Four pairs of branchiae; three smooth and one pennate; small paleae present. Glandular ridges absent. Thirteen thoracic uncinigers. Sabellides Milne Edwards in Malmgren 1966, Sabe lia octocirrata Sars 1835; 8 species. AMPHARETINAE, Four pairs of smooth branchiae; small paleae reported in some species. Glandular ridges and abdominal notopodial rudiments absent. Eleven thoracic uncinigers. Notosetae absent in segment 4. Buccal tentacles papillose. Samytha Malmgren 1866, Sabellides sexcirrata Sars 1856; 7 species. AMPHARETINAE. Three pair? of smooth branchiae; paleae absent. Glandular ridges absent; abdominal notopodial rudiments present. Fourteen thoracic uncinigers. Samythella Verrill 1873a, S. elongata Verrill 1873a; 6 species.
128
Science Series 28
AMPHARETINAE. Three pairs of smooth or flanged branchiae. Paleae absent. Glandular ridges absent; abdominal notopodial rudiments absent. Twelve thoracic uncinigers. Circle of anal papillae. Samythopsis Mclntosh 1885, S. grbet Mclntosh 1885; only species. AMPHARETINAE. Three pairs of smooth branchiae; paleae absent. Glandular ridges and abdominal notopodial rudiments present. Fourteen thoracic uncinigers. Schistocomus Chamberlin 1919a, 5. hiltoni Chamberlin 1919a; 3 species. AMPHARETINAE. Four pairs of branchiae, one smooth and three lamellate; paleae absent. Glandular ridges absent; abdominal notopodial rudiments present. Twelve thoracic uncinigers. Two pairs of anal cirri. Sosane Malmgren 1866, S. sulcata Malmgren 1866; 5 species. AMPHARETINAE. Four pairs of smooth branchiae; paleae present. Glandular ridges absent; abdominal notopodial rudiments present. Twelve thoracic uncinigers. Third from last thoracic notopodia elevated and with modified setae. Sosanella Hartman 1965, S. apalea Hartman 1965; only species. AMPHARETINAE. Three pairs of smooth branchiae; paleae absent. Glandular ridges and abdominal notopodial rudiments absent. Thirteen thoracic uncinigers. Notopodia in setiger 13 (third from last thoracic notopodium) elevated with hirsute setae. Sosanides Hartmann-Schrder 1965, S. glandularis Hartmann-Schrder 1965; only species. AMPHARETINAE. Four pairs of smooth branchiae; paleae present. Abdominal notopodial rudiments absent. Twelve thoracic uncinigers. Setiger 11 with modified notosetae. Sosanopsis Kessle 1917, S. wireni Hessle 1917; 2 species. AMPHARETINAE. Four pairs of smooth branchiae; paleae absent. Glandular ridges absent; abdominal noto-
podial rudiments present; 12 thoracic uncinigers. Notosetae of last thoracic setiger modied, Weddellia Hartman 1967, W. profimda Hartman 1967; only species. AMPHARETINAE. Three pairs of smooth branchiae on three successive segments. Segment 3 with capillary setae, but without paleal modifications. Glandular ridges absent; abdominal parapodia with long dorsal cirri. Fifteen thoracic uncinigers.
Invalid Genera Aryandes Kinberg 1867b, indeterminable Branchiosabella Claparde 1863, set Ampharete Crossostoma Gosse 1855, sec Amphicteis Eusamytha Hartman 1967, see Eusamythella Heterobranchus Wagner 1885, ses Sabellides Melinnides We sen berg-Lund 1950, see Melinnexis Annenkova, 1931 Melinnopsis Day 1955, see Melinnopsides Microsamytha Augener 1928a, see Alkmaria Pseudosabellides Berkeley and Berkeley 1943, see Asabellides Rytocephalus Quatrefages 1866, indeterminable
FAMILY TEREBELLIDAE MALMGREN
1867
Body in two regions; anterior region with biramous parapodia and posterior region with neuropodia only. Piostomium a simple fold. Branchiae, when present, include one to three pairs on the first segments, associated distinctly with separate segments. Uncini usually with a large main fang and a crest of smaller teeth. Terebellids are among the most common shallowwater polychaetes and are found in all environments. The usually numerous buccal tentacles cannot be fully retracted into the mouth. They usually are grooved and used in selective deposit-feeding on the surface. Other forms may stretch the buccal tentacles into the water and capture particles from the water.
Key to Genera la. lb. 2a (lb). 2b (lb). 3a (2b). 3b (2b). 4a (la). 4b (la). 5a (3a). A large scoop-shaped or conical proboscis present ARTACAMINAE 4 Proboscis absent 2 Thoracic uncini in double or alternating rows in at least some setigers .... AMPHTTRITINAE ... 14 Thoracic uncini in single rows in all setigers 3 Branchiae present THELEPINAE 5 Branchiae absent POLYCIRRINAE 10 I>roboscis papillated, 17 thoracic setigers Artacama Proboscis grooved; 15 thoracic setigers Artacamella First notosetae on first postbranchial segment Pseudampharete
7976 5b (3a). 6a (5b). 6b (5b). 7a (6b). 7b (6b). 8a (7a). 8b (7a). 9a (7b). 9b (7b). 10a (3b). 10b (3b). I la ( 10b). II b (10b). 12a (1 la). 12b (1 la). 13a (1 lb). 13b (lib). 14a (2a). 14b (2a). 15a (14a). 15b ( 14a). 16a (15a). 16b ( 15a). 17a (16a). 17b (16a). 18a (17b). 18b (17b). 19a (16b). 19b (16b). 20a (19b). 20b (19b). 1\2L (15b). 21b (15b). 22a (14b). 22b (14b). 23a (22a). 23b (22a). 24a (23b). 24b (23b). 25a (24b). 25b (24b). 26a (25a). 26b (25a). 27a (26b). 27b (26b). 28a (24b). 28b (24b). 29a (28a). 29b (28a). 30a (28b). 30b (28b). 31a (22b). 31b (22b). 32a (31a). 32b (31a). 33a (32b).
129
First notosetae on one of the branchial segments 6 First notosetae on the first branchial segment Streblosoma First notosetae on the second branchial segment 7 Lateral lappets present on one or a few anterior segments 8 Lateral lappets absent 9 Uncini from the fifth segment Euthelepus Uncini from the first abdominal segment Telothelepus Uncini from seliger 9 Parathelepus Uncini from setiger 2 Tkelepus Setae completely absent Hauchiella At least some setae present 11 Neurosetae absent 12 Neurosetae present 13 Thoracic notopodia vascularized and in part furcate or branched Enoplobranchus Thoracic notopodia may be vascularized, but never furcate or branched Lysilla Neurosetae short-handled uncini Polycirrus Neurosetae long-handled spines Amaeana Branchiae absent 15 Branchiae present 22 Seventeen or fewer thoracic setigers 16 More than 20 thoracic setigers 21 All notosetae smooth-tipped 17 At least some notosetae with denticulated tips 19 Lateral lappets absent; 17 thoracic setigers Laphania Lateral lappets present on one or a few anterior segments; 16 thoracic setigers 18 Third segment with transverse ridge across dorsum Leaena Third segment without transverse ridge Stschapovella Neurosetae from setiger 3; some notosetae smooth Proclea Neurosetae fiom setiger 2; all notosetae denticulated 20 Fifteen thoracic setigers; notosetae finely denticulated Lanassa Fourteen thoracic sefigers; some notosetae distinctly pectinate Phisidia All notosetae tapering, distally smooth-tipped Baffinia Some notosetae subdistally inflated, distally denticulated Spinosphaera Notosetae with marginally serrated tips 23 Notosetae with smooth tips 31 Three anterior segments with large lateral lappets to which the branchiae are attached . Colymmatops Lateral lappets present or absent, but never directly associated with the branchiae 24 Uncini from third thoracic setiger Neoleprea Uncini from second thoracic setiger 25 Sixteen or 17 thoracic setigers 26 Eighteen or more thoracic setigers 28 A single pair of branchiae; 16 thoracic setigers Spiroverma Two or three pairs of branchiae; 17 thoracic setigers 27 Lateral lappets present; branchiae stalked and branched Neoamphitrite Lateral lappet absent; branchiae sessile filaments Amphitrite Branchiae attached on segments 2-4 29 Branchiae attached on another combination of segments 30 Branchiae arbwrescent from the base Terebella Branchiae distinctly stalked Amphitritides Branchiae on segments 1, 2 and 5 Polymniella Branchiae on segments 3, 7 and 13 Terebellobranchia Anterior uncini long-handled 32 All uncini short-handled 35 Notosetae from segment 5, uncini from segment 6 Opisthopista Notosetae from segment 4, uncini from segment 5 33 Branchiae smooth, ampharetinlike Eupistella
130 33b (32b). 34a (33b). 34b (33b). 35a (31b). 35b (31b). 36a (35b). 36b (35b). 37a (36a). 37b (36a). 38a (37b). 38b (37b). 39a (38b). 39b (38b). 40a (39b). 40b (39b). 41a (4b). 41b (40b). 42a (41b). 42b (41b). 43a (42b). 43b (42b). 44a (36b). 44b (36b). 45a (44b). 45b (44b). 46a (45b). 46b (45b). 47a (46b). 47b (46b). 48a (47b). 48b (47b).
Science Series 28
Branchiae branched 34 Lateral lappets at least on segments 2 and 4 Pista Lateral lappets limited to segment 3 Lanicides Ventrum with large anteriorly opening glandular folds on segments 3-5 Paraxionice Ventrum with glandular scutes or smooth 36 Lateral lappets present on one or a few anterior segments 37 Lateral lappets inconspicuous or absent 44 Sixteen thoracic setigers; branchiae very long-shafted Axionice Seventeen or 18 thoracic setigers; branchiae short-shafted or sessile 38 Eighteen thoracic setigers Melinella Seventeen thoracic setigers 39 First four segments flattened with large lateral lappets forming an anterior plaque; one pair of branchiae Scionella First four segments not modified into a flattened plaque; lateral lappets varying in size; two or three pairs of branchiae 40 Branchiae tufts of sessile filaments Thelepides Branchiae branching off one or a few stems 41 Buccal lateral lappets form a crest across the dorsum Paralanice No dorsal crest across the buccal segment 42 Uncini with teeth in a single row Loimia Uncini with teeth in two or more rows 43 Small lateral lappets on segments 2 and 3; tube opening unadorned Eupofymnia Large lateral lappets on segments 2 and 4; tube opening with fan-shaped frills Lanice Thirteen thoracic setigers Ramex At least 15 thoracic setigers 45 First notosetae on first branchial segment Naneva First notosetae on last branchial segment or from one of the first postbranchial segments 46 Uncini present from first setiger Reieterebeila Uncini present from second setiger 47 Uncini in an open circle in posterior thoracal setigers Terebellanice Uncini arranged back to back in posterior thoracal setigers 48 Two pairs of branchiae; thorax with 15 to 40 setigers Nicolea Three pairs of branchiae; thorax with 17 setigers Scionides
Generic Definitions Amaeana Hartman 1959, Polycirrus trilobatus Sars 1863- 5 species POLYCIRRINAE. Tentacular lobe trifoil-shaped. Ten to 13 thoracic segments; notosetae from segmem 3. Uncini absent; abdominal neurosetae long-shafted spines. Amphitrite O.F. Mller \11\,A. cirrata O.F. Mller 1771; 17 species. AMPHITRITINAE. Eyes rarely present. Three pairs of sessile branchiae from segment 2; nephridial papillae on segment 3 and fi-om segment 6. Lateral lappets absent. Notosetae from fourth segment; distally serrated; 13 to 25 thoracic setigers. Amphitritides Augener 1922, Terebella gracilis Grube 1860; 3 species. AMPHITRITINAE. Two pairs of branching, stalked branchiae from segment 2. Lateral lappets absent. Serrated notosetae present from segment 4; uncini face to face in posterior thoracic setigers.
Artacama Malmgren 1866, A. proboscidea Malmgren ^^^^' ^ ^P^*^'**ARTACAMINAE. Three pairs of filamentous ''^^'^Wae on segments 2-4. Lateral lappets absent. Sevemeen thoracic segers; notosetae distally smooth. P^P'" proboscis attached ventrally on the peris"""*""' Artacamella Hartman 1955, A. hancocki Hartman 1955; 2 species. ARTACAMINAE. Three pairs of smooth, ampharetinlike branchiae on segments 2-4. Fifteen thocic setigers; uncini in all thoracic setigers; each uncinus long-shafted. Grooved, boat-shaped proboscis attached ventrally on the peristomium. Axionice Malmgren 1866, Terebella flexuosa Grube 1860; 8 species. AMPHITRITINAE. Two or three pairs of longshafted, branched branchiae. Sixteen thoracic setigers; lateral lappets present. Notosetae distally smooth; all uncini short-handled.
}976
Baffinia Wesenberg-Lund 1950, B. multisetosa Wesenherg-Lund 1950; only species. AMPHITRITINAE. Branchiae absent. Notsetae present from third segment to the end of the body (more than 70 segments); uncini present from second setiger; uniserial in first eight uncinigers, then biserial and finally uniserial in last 30-35 setigers. Capillary setae distally smooth. Bathya Saint-Joseph 1894, Leaena abyssorum McIntosh 1885; 3 species. AMPHURITINAE. Branchiae absent. Uncini with short handles; crested; capillary distally smooth, resembles Proclea in setal structures. Incompletely described. Cotymmatops Peters 1854, C. granulatus Peters 1854; only species, AMPHITRITINAE. Three first segments with large lateral lappets to which are attached branchiae. Thirteen or 14 thoracic setigers. Notosetae distally serrated. Incompletely described. Enoplobranckus Webster 1879, Chaetobranchus sangunea Verrill 1873b; only species. POLYCIRRINAE. Notopodial lobes prolonged, vascularized and in part furcate or branched. Uncini absent; notosetae spinose capillaries, with usually one seta much longer than the others. Eupistella Chamberlin 1919c, Eupista darwini McIntosh 1885; 4 species. AMPHITRITINAE. Two pairs of smooth, ampharetinlike branchiae. Seventeen thoracic setigers; some anterior uncini with prolonged shafts, notosetae distally smooth. Eupolymnia Verrill 1900, Amphitrite nesidensis delle Chiaje 1828; 12 species. AMPHITRITINAE. Three pairs of branching branchiae; lateral lappets on segments 2-3. Smooth-tipped notosetae from segment 4; 17 thoracic setigers. Euthelepus Mclntosh 1885, E. setubalensis Mclntosh 1885; 6 species. THELEPINAE. Branchiae on segments 2-4, sometimes as single filaments only. Lateral lappets present. Notosetae from second branchial segment; present on 20 segments. Uncini first present from first postbranchial segment. Hauchiella Levinsen 1893, Polycirrus tribullata Mclntosh 1869; only species. POLYCIRRINAE. Thorax of ten segments; usually about 70 segments in all. All setae absent. Lanassa Malmgren 1866, L. nordenskioldi Malmgren 1866; 7 species. AMPHITRITINAE. Branchiae absent. Lateral lappets sometimes present. Fifteen thoracic setigers. Notosetae present from segment 4, denticulated tips.
Lanice Malmgren 1866, Nereis conchilega Pallas 1766; 8 species. AMPHITRITINAE. Three pairs of branched branchiae. Lateral lappets present. Seventeen thoracic setigers. Smooth-tipped notosetae from segment 4; uncini back to back in posterior thoracic setigers. Tube with branched fine-meshed fan attached to opening. Lanicides Hessle 1917, Terebella (Phyzelia) bilobata Grube 1877; 3 species. AMPHITRITINAE. Two pairs of branched branchiae; lateral lappets present. Smooth-tipped notosetae present from segment 4; long-shafted uncini present from segment 5. Laphania Malmgren 1866, L. boecki Malmgren 1866; only species. AMPHITRITINAE. Branchiae absent. Lateral lappets absent. Seventeen thoracic setigers; notosetae distally smooth. Uncini present from setiger 7. Leaena Malmgren 1866, Terebella abranchiata Sars 1865; 10 species. AMPHITRITINAE. Branchiae absent. Lateral lappets present; third segment with transverse ridge across dorsum. Smooth-tipped notosetae present from segment 4. Sixteen thoracic setigers. Loimia Malmgren 1866, Terebella medusa Savigny 1818; 16 species. AMPHITRITINAE. Three pairs of branched branchiae. Lateral lappets present. Seventeen thoracic setigers; notosetae smooth-tipped. Uncini with all teeth in a single row (pectinate). Lysilla Malmgren 1866, L. loveni Malmgren 1866; 10 species. POLYCIRRINAE. Six to 12 thoracic segments; notosetae from segment 3. Neurosetae completely absent. Melinetla Mclntosh 1914, A/, macduffi Mclntosh 1914; only species. AMPHITRITINAE. One pair of branched branchiae; 18 thoracic setigers, all with uncini. Lateral lappets present. Naneva Chamberlin 1919a, A^. hspera Chamberlin 1919a; only species. AMPHITRITINAE. Two pairs of dendritically branched branchiae from segment 2; smooth-tipped notosetae present from first branchial segment. Twentyseven thoracic setigers. Lateral lappets absent. Uncini present in double-rows in most thoracic segments. Neoamphitrite Hessle 1917, Amphitrite affinis Malmgren 1866; II species. AMPHITRITINAE. Three pairs of branched branchiae; lateral lappets present. Seventeen thoracic- setigers with distally serrated notosetae. Nephridial papillae present from segment 3.
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Neoleprea Kessle 1917, Leprea streptochaeta Ehlers 1897; 5 species. AMPHITRmNAE. Two or three pairs of branched branchiae; lateral lappets absent. Notosetae first present from segment 3; some smooth, some distally denticulated. Seventeen-40 thoracic setigers. Nephridial papillae present on segments 3-9. Nicolea Malmgren 1866, Terebella zostericola rstcd 1844; 22 species. AMPHITRmNAE. Two pairs of branched branchiae. Lateral lappets absent. Smooth-tipped notosetae present from segment 4; 15-40 thoracic setigers. Uncini arranged back to back in posterior thoracic segments. Opisthopista Caullery 1944, O. sibogae Caullery 1944; only species. AMPHITRITINAE, Two pairs of branched branchiae; lateral lappets present on at least segments 2 and 4. First notosetae in segment 5 and first uncini in segment 6. Anterior uncini long-shafted. Paralanice Caullery 1944, P. timorensis Caullery 1944; only species. AMPHITRITINAE. Three pairs of branched branchiae; large lateral buccal lappets connected across dorsum with a crest; lateral lappets also on segments 2 and 3. Seventeen thoracic setigers; smooth-tipped capillaries from segment 4. Parathelepus Caullery 1915, Thelepides collaris Southem 1914; only species. THELEPINAE. Three pairs of branchiae; lateral lappets absent. Notosetae from second branchial segment; uncini from setiger 9. Paraxionice Fauchald 1972, P. artifex Fauchald 1972; only species. AMPHITRITINAE. One pair of branched branchiae with double bases on segments 3 and 4. Sixteen thoracic setigers; notosetae distally smooth. Ventral part of segments 3-5 covered by a large glandular apparatus that opens anteriorly on segment 3 in two trumpetshaped openings. Phisidia Saint-Joseph 1894, Leaena oculata Langeriians 1880; 3 species. AMPHITRITINAE. Branchiae absent; lateral lappets absent. Notosetae distally denticulate; longer finely so, the shorter coarse, with pectinate appearance. Uncini from setiger 2; 14 thoracic setigers. Pista Malmgren 1866, Amphitrite cristata O.F, Mller 1776; 40 species. AMPHITRmNAE. Two pairs of stalked branched branchiae; lateral lappets large, on segments 2 and 4 at least. Smooth-tipped notosetae from segment 4; long-handled uncini present in anterior setigers. Fifteen to 24 thoracic setigers.
Polycirrus Grube 1850, P. medusa Grube 1850; 39 species. POLYCIRRINAE. Thorax with a variable number of setigers; notosetae present from segment 3. Uncini first present from segments 7-18. Notosetae distally smooth or serrated. Polymniella Verrill 1900, P. aurantiaca Verrill 1900; only species. AMPHITRITINAE, Three pairs of branched branchiae on segments 1, 2 and 5. Notosetae from segment 1, marginally dentate; uncini from segment 2. Twentytwo or more thoracic segments. Proclea Saint-Joseph 1894, Leaena graffii Langerhans 1880; 3 species. AMPHITRITINAE. Branchiae absent. Lateral lappets present. Sixteen thoracic setigers; notosetae distally either dentate or smooth. Uncini from setiger 3, Pseudatnpharete Hartmann-Schrder 1960b, P. tentaculata Hartmann-Schrder 1960b; only species. THELEPINAE. Two pairs of sessile branchial filaments on large bosses on first and second segment. Notosetae first present from first postbranchial segment; uncini from setiger 7. Ramex Hartman 1944b, R. californiensis Hartman 1944b; only species. AMPHITRmNAE. One pair of branched branchiae on second segment. Notosetae from segment 4; thorax with 13 setigers; notosetae distally smooth. Reteterebella Hartman 1963b, R. queenslandia Hartman 1963b; only species, AMPHITRmNAE. Three pairs of branched branchiae; lateral lappets inconspicuous. Sixteen thoracic setigers; notosetae from segment 4; distally smooth. Uncini present from first setiger. Scionella Moore 1903, S. japnica Moore 1903; 4 species. AMPHITRITINAE. One pair of branchiae on segment 4. Seventeen thoracic setigers; notosetae first present on segment 4; notosetae distally smooth. Four first segments flattened dorsoventrally with very large longitudinally oriented lateral lappets, forming a large oblique plaque at the anterior end. Scionides Chamberlin 1919b, Terebella reticulata Ehlers 1887; 2 species. AMPHITRITINAE. Three pairs of branched branchiae; seventeen thoracic setigers; notosetae from segment 4; notosetae distally smooth. Uncini arranged back to back in posterior thoracic segments. Spinosphaera Kessle 1917, S. pacifica Kessle 1917; 2 species. AMPHITRmNAE, Branchiae absent. Twenty-three or more thoracic setigers; uncini present from setiger 2,
1976
Notosetae denticulate; the longer ones with hispid swellings subdistally. Lateral lappets absent. Spiroverma Uchida 1968, S. ononokomachii Uchida 1968; only species. AMPHITRITINAE. One pair of sessile branchiae, each with maximally eight filaments on segment 2. Sixteen thoracic setigers; notosetae marginally serrate. Body strongly spiralled. Streblosoma Sars 1872, Grymaea bairdi Malmgren 1866; 20 species. THELEPINAE. Three pairs of sessile branchiae on segments 2-4 (may be absent). Notosetae from first branchial segment (segment 2). Uncini from segment 5. Stschapovella Levenstein 1957, S. tatjanae Levenstein 1957; only species. AMPHITRITINAE. Branchiae absent; lateral lappets present. Smooth-tipped notosetae from segment 4; uncini from segment 5. Sixteen thoracic setigers. Notosetae finely capillary rather than limbate. All nephridia free from one another. Telotheiepus Day 1955, T. capensis Day 1955; only species. THELEPINAE. Two or three pairs of branchiae from segment 2. Notosetae from second branchial segment (segment 3). Lateral lappets present. Fifteen thoracic setigers. Tentacular lobe very large and frilly. Neuiosetae absent on thorax, uncini present on abdomen. Terebella Linnaeus 1767, T. lapidaria Linnaeus 1767; 28 species. AMPHITRITINAE. Two or three pairs of branched branchiae; lateral lappets absent. Thorax with variable, usually large, number of setigers; notosetae from segment 4; distally serrated. Uncini face to face in posterior thoracic segments. Terebellanice Hartmann-Schrder 1962b, T. laeviseta Hartmann-Schrder 1962b; only species. AMPHITRITINAE. Two pairs of branchiae from third segment; lateral lappets absent. Notosetae smoothtipped. Uncini in an open circle on posterior thoracic segments. Terebellobranchia Day 1951, T. natalensis Day 1951; 2 species. AMPHITRITINAE. Three pairs of branched branchiae on segments 3, 7 and 13. Distally serrated notosetae present from segment 4; more than 19 thoracic setigers present. Thelepides Gravier 1911a, T. koehleri Gravier 1911a; 3 species. AMPHITRITINAE. Three pairs of filiform branchiae; lateral lappets present. Smooth-tipped notosetae fi-om segment 3; 17 thoracic setigers present.
Thelepus Leuckart 1849, Amphitrite cincinmta Fabricius 1780; 32 species. THELEPINAE. Sessile, filiform branchiae on seg ments 2-4; smooth-tipped notosetae present from second branchial segment (segment 3). Uncini from segment 5.
Taxonomic Notes Day (1967) altered the definition of the sub-families to include the abranchiate members of AMPHITRITINAE among the POLYCIRRINAE. This change appears unfortunate, in that these genera resemble branchiate members of the AMPHITRITINAE very closely in setal structures as well as in the structure of the anterior end. The treatment here reflects this view. The genus Pseudampharete has been included among the THELEPINAE since it is branchiate and has the uncini in single rows in all thoracic segments. It further resembles members of this subfamily in that the branchiae are sessile filaments. However, this latter feature may also be present among members of the AMPHITRITINAE. The placement must be considered temporary. The genus Batkya Saint-Joseph 1894, listed above in the definitions has not been included in the key. It belongs in the abranchiate gorup of AMPHITRITINAE, and is very poorly known. The genus Pseudothelepus Augener 1918 is considered here a synonym of Streblosoma, as suggested by Day (1967). Some of the genera are difficult to separate from related forms; no revision was attempted on this occasion.
Invalid Genera Amaea Malmgren 1866, ss&Amaeana Amphiro Montagu 1808, se& Amphitrite Amphitritoides Costa 1862, ste Eupolymnia Amphytrite Renier 1804, indeterminable Anisocirrus Gravier 1905a, see Polycirrus Aphlebina Claparde 1864, see Polycirrus Apneuma Quatrefages 1866, see Polycirrus Athelepus Chamberhn 1919c, NOMEN NUDUM Chaetobranchus Verrill 1873b, see Enoplobranckus Cyaxares Kinberg 1867b, see Polycirrus Dejoces Kinberg 1867b, see Polycirrus Dendrobranchus Wagner 1885, indeterminable Dendrophora Grube 1870a, see Pista Ehlersiella Mclntosh 1885, indeterminable Ereutho Malmgren 1866, see Polycirrus Eugrymaea Verrill 19(X), see Streblosoma Eupista Mclntosh 1885, see Eupistella Euscione Chamberlin 1919c, seeAxionice Grymaea Malmgren 1866, see Streblosoma Heterophenacia Quatrefages 1866, see Thelepus
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Heterophyselia Quatrefages 1866, see Terebella Heteroterebella Quatrefages 1866, see Terebella dalia Quatrefages 1866, &ee Pista Laphaniella Malm 1874, setLanassa LepreaMa\mgren 1866, stc Terebella Leucariste Malmgren 1866, se^ Pofycirrus Lumara Stimpson 1854, see Thelepus Neottis Malmgren 1866, see Thelepus Odysseus Kinberg 1867b, indetenninable Otanes Kinberg 1867b, see Pista Pallonia Costa 1862, see Eupolymnia Phenacia Quatrefages 1866, see Thelepus Pherea Saint-Joseph 1894, sttLanassa Phyzelia Schmarda 1861, seeNicolea Pofymnia Malmgren 1867, see Eupolymnia Protothelepus Verrill 1900, see Euthelepus Schmardanella Mclntosh 1885, see Terebella Scione Malmgren 1866, seeAxionice Scionopsis Verrill 1873b, see Pista Solowetia Ssolowiew 1899, seeProclea Thelepella Chamberlin 1919c, seeNicolea Thelephusa Verrill 1871, see Thelepus Tkelepides Southern 1914, see Parathelepus Thelepodopsis Sars 1872, see Thelepus Torquea Leidy 1855, see Pofycirrus
Uncinochaeta Quatrefages 1866, indeterminabie Venusia Johnston 1865, see Thelepus Wartelia Giard 1878, seeLanice
FAMILY TRICHOBRANCHIDAE MALMGREN
1866
Body separated into two regions; anterior region with biramous parapodia, posterior region with neuropodia only. Prostomium a large hood over the mouth. Maximally four pairs of branchiae present on anterior setigers. Anterior neuropodia with long-shafted hooks; abdomen with densely crested uncini, Notosetae capillary. The trichobranchids are related closely to the terebellids and frequently are included among them as a subfamily (Day 1967). They differ rather strikingly in body-proportions, in that they tend to be slenderer and more muscular than the terebellids. The large folded prostomium also gives them a characteristic appearance. One species, Terebellides stroemi, has been reported from worldwide areas, and is apparently nearly everywhere, an important member of soft-bottom environments. Members of the family appear to be most common in cold-water soft bottoms and may be represented best in deep water. Key to Genera
la. lb. 2a (la). 2b (la). 3a (lb). 3b (lb). 4a (3a). 4b (3a). 5a (3b). 5b (3b). 6a (5b). 6b (5b).
A single middorsaUy attached branchia present At least a pair of branchiae present Branchia a single tapering, digitate projection Branchia stalked with four lobes; each lobe with numerous flat branchial lamellae Two or three pairs of branchiae present Four pairs of branchiae present , Two pairs of branchiae, 17 thoracic setigers Three pairs of branchiae; 15 thoracic setigers Each branchia with pectinate branchial lamellae Each branchia simple and digitate or rosette-shaped All branchiae simple and tapering Fourth pair of branchiae sessile rosettes Generic Definitions
2 3 Unobranchus Terebellides 4 5 Filibranchus Trichobranchus Ampharetides 6 Octobranchus Novobranchus
Ampharetides Ehlers 1913, A. vanhoeffeni Ehlers 1913; only species. Seventeen thoracic setigers; uncini present on nine last thoracic setigers and on abdomen. Four pairs of branchiae on first setiger; each with pectinate branchial lamellae. Filibranchus Malm 1874, F. roseus Malm 1874; only species. Fifteen thoracic setigers; all with uncini. Two pairs of smooth brapchiae on the second and third segment (presetal branchiae).
Novobranchus Berkeley and Berkeley 1954, N. pacificus Berkeley and Berkeley 1954; only species. Sixteen thoracic setigers; uncini present ftom setiger 4. Four pairs of branchiae on segments 2-5; first three pairs lanceolate; last pair a frilled rosette. Octobranchus Marion and Bobretzky 1875, Terebella lingulata Grube 1863; 4 species. Sixteen thoracic setigers; first setiger is segment 3; first uncini on setiger 4. Four pairs of similar lanceolate branchiae. Terebellides Sars 1835, T. stroemi Sais 1835; 14 species.
1976
/J5 Invalid Genera
Eighteen thoracic setigers; first seliger is segment 3; uncini present from seliger 6. Single dorsal branchia with four lamellate branchial lobes. Trichobranchus Malmgren 1866, T. glacialis Malmgren 1866; 5 species. Fifteen thoracic setigers; first setae on segment 6; all thoracic setigers with uncini. Three pairs of simple, lanceolate branchiae. Vnobranchus Hartman 1965, U. abyssalis Hartman 1965; only species. Twelve thoracic setigers; first setae on segment 3; uncini first present on setiger 4. A single, large lanceolate branchia present mid-dorsally on segment 2.
Aponobranchus Gravier 1905a, see Terebellides Corephorus Grube 1846, see Terebellides
FAMILY BOGUEIDAE HARTMAN AND
FAUCH ALD
1971
Body slender and cylindrical with few segments; prostomium rounded without appendages. Parapodia biramous with long slender, in part spinose notosetae and short-handled, avicular crested uncini in the neuropodia, in part in double rows. The bogueids, known for two genera, have setae resembling the terebellid uncini, but lack completely anterior appendages. They are known from a few locations in the western Atlantic Ocean only.
Key to Genera la. lb. First three setigers without neurosetae First four setigers without neurosetae Boguea Boguella
Generic Definitions Boguea Hartman 1945, B. enigmtica Hartman 1948; only species. Two anterior asetigerous segments; three first setigers without neurosetae. Boguella Hartman and Fauchald 1971, . ornato Hartman and Fauchald 1971; only species. One anterior asetigerous segment; four first setigers without neurosetae.
ORDER SABELLIDA Prostomium reduced, fused with the peristomium which usually forms a large tentacular crown; setae include thoracic notopodial limbate and geniculate kinds and neuropodial uncini; setal positions reversed in the abdomen.
FAMILY SABELLIDAE MALMGREN
1867
Body cylindrical with a thorax of few setigers and abdomen with few to many. Uncini crested or with teeth in several rows; long- or short-handled. Tube present in most species, made of varying material, but never calcareous. Sabellids characteristically have nearly smooth appearing bodies, cylindrical and tapering posteriorly, with large, often maroon or red-colored tentacular crowns. Most of the forms, especially the larger ones,
are strictly sessile and never leave their tubes; the smaller forms, such as species of Fabricia and allied genera, are capable of moving around. Most of the larger forms are associated with shallow water; smaller species are common in deep sea collections. The major characteristics used to identify the sabellids, include the presence or absence of companion setae to the neuropodial uncini in the thorax; these also have been called pennoned setae or pick-ax setae. The neutral term companion seta is preferred here. They occur in an anterior row, in front of the uncini they accompany and usually are small and deeply imbedded in the epidermal tissues. They are more easily seen by the reflection they give off under a stereo microscope, than in microscopic preparation under the compound microscope. The structure of the tentacular crown such as the number of radioli, and the presence of small, external appendages called sty lodes, are important. One of the key features in the group lies in the structure of the thoracic uncini. These may be acicular, by which is meant that the crested head, with one large tooth and several smaller ones, is supported by a gently curved, often nearly straight shaft. By avicular uncini are meant uncini that are essentially Z-shaped (sometimes called swan-shaped) with the small crested head at the top of the Z, and the shaft sharply bent. The shafts of these Z-shaped uncini may be short or long. In the key below, attempts have been made to avoid the more confusing part of the terminology, but a complete avoidance of this terminology has not been possible.
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FIGURE 37. (A), Family SABELLIDAE, Chone sp., off Santa Catalina Island, 50 m, 18x; (B), Famy SABELLONGIDAE, Sabellonga disjoncta, anterior end in right ventrolateral view, modified after Hartman, 1969, 33x; (C), posterior end of the above, 33x; (D), Family CAOBANGI)AE, Caobangia abbotti, after Jones, 1974, about 17x; (E), right lateral view of the above, about I7x.
Key to Genera lalb. 2a (lb). 2b (lb). 3a (2a). 3b (2a). MYXICOLINAE Myxicola Abdominal uncini in short, discrete tori 2 Thoracic uncini with long, gently curved shafts (acicular) companion setae always absent FABRICINAE 3 Thoracic uncini with short or long, but always strongly bent shafts (avicular); companion setae present in some forms SABELLINAE 15 Abdomen with two or three setigers 4 Abdomen with four or more setigers 9 Abdominal uncini form nearly complete cinctures around the body
]76 4a (3a). 4b (3a). 5a (4b). 5b (4b). 6a (5b). 6b (5b). 7a (6b), 7b (6b). 8a (7b). 8b (7b). 9a (3b). 9b (3b). 10a (9a). 10b (9a). 11 a (9b). 1 lb (9b). 12a (1 lb). 12b (1 lb). 13a (12b). 13b ( 12b). 14a (13b). 14b (13b). 15a (2b). 15b (2b). 16a (15a). 16b (15a). 17a (16a). 17b (16a). 18a (16b). 18b (16b). 19a (18a). 19b (18a). 20a (18b). 20b (18b). 21a (15b). 21b (15b). 22a (21a). 22b (21a). 23a (22b). 23b (22b). 24a (23a). 24b (23a). 25a (23b). 25b (23b). 26a (2lb), 26b (21b). 27a (26b). 27b (26b). 28a (27b). 28b (27b). 29a {28b). 29b (28b). 30a (29b). 30b (29b). 31a (30b).
^7
N,
Radiles partially united by a membrane Monroika Radiles free to the base 5 With an extensive lower lip, curved up under the weakly bilobed prostomium Pseudofabricia Without a distinct lower lip; prostomium anteriorly rounded 6 Abdominal uncini short-handled Fabriciola Abdominal uncini long-handled 1 Without ventral vascularized filaments Fabricia With ventral vascularized filaments in the tentacular crown 8 Ventral vascularized filaments branched; three pairs of radioli Augeneriella Ventral vascularized filaments smooth; two pairs of radioli Mamymkia Posterior end distinctly modified, with fused segments or greatly expanded pygidium 10 Posterior end not modified 11 Seven last setigers covered dorsaily by the large, ring-shaped pygidium Euchonella Last several setigers flanged laterally to form a spoon-shaped dorsal cavity Euchone Radiles united by a membrane Chone Radiles free to the base ' 12 Abdominal uncini with a large main fang and a crest of smaller teeth Jasmineira Abdominal uncini with major teeth in two or more rows 13 More than ten pairs of radioli in the tentacular crown Dialychone Maximally five pairs of radioli in the tentacular crown 14 Radiles externally distinctly ridged Oriopsis Radiles externally rounded Desdemom Thoracic neuropodial companion setae absent 16 Thoracic neuropodial companion setae present 21 Radioli with external stylodes 17 Radiles without external stylodes 18 Four thoracic setigers, external stylodes very small Pseudobranchiomma Eight thoracic setigers; external stylodes well developed Branchiomma Thoracic spatulate notosetae present 19 Thoracic spatulate notosetae absent 20 Thoracic uncini with gently curved, long handles Fabrisabella Thoracic uncini with sharply bent handles Laonome Collar reduced, radioli basally united by a web Euratella Collar well developed; radioli free to the base Sabellastarte Tentacular crown with radioli arranged in one or two spirals 22 Tentacular crown with radioli in two semicircles 26 Posterior thoracic notopodia with lancet-shaped, transversely striated setae Bispira Lancet-shaped notopodial setae absent 23 Collar two-lobed 24 Collar four-lobed 25 Anterior ventrura irregularly rugose Distylidia Anterior ventrum smooth, apart from the glandular fields Demonax One half of tentacular crown very much larger than the other; spatulate setae absent .. Spirographis Both parts of the tentacular crown similar in size; spatulate setae present Eudistylia Large subdistal compound eyes on a few dorsal radioli Megalomma Eyes, if present, smaller and scattered over most of the radioli 27 Collar reduced or absent, a triangular ventral projection at base of tentacular crown present Amphigiena Collar well developed; triangular ventral projection absent 28 Notosetae of the collar-segment in a long row Hypsicomus Notosetae of the collar-segment in a tuft 29 Radioli dichotomously branching Schiiobranchia Radioli not divided 30 Spatulate thoracic notosetae absent Sabella Spatulate thoracic notosetae present 31 Thoracic uncini at least in part nearly straight and long-handled 32
38 31b 32a 32b 33a 33b (30b). (31a). (31a). (31b). (3Ib).
NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY Thoracic uncini sharply bent, long- or short-handled Abdominal uncini long-handied and nearly straight Abdominal uncini short-handled and strongly bent Thoracic uncini long-handied Thoracic uncini short-handled
Science Series 28 33 Potaspina Potamethus Panousea Potamilla
Generic Definitions Ampkiglena Claparde 1864, Amphicora mediterrnea Ley dig 1851; 3 species. SABELLINAE. Radiles in semi-circles; external stylodes, eyes and webbing absent. Collar absent; triangular ventral projection present at base of tentacular crown. Approximately eight thoracic setigers present; abdominal uncini strongly bent, crested. Augeneriella Banse 1957,^. hummelincki Banse 1958; 4 species. FABRICINAE. Three pairs of radioli, branchial heart present; collar dorsally low, ventrally higher. Forked ventral vascularized filament present. Eight thoracic and three abdominal setigers; abdominal uncini long-handled. Bispira Kryer 1856, Amphitrite volutacornis Montagu 1804; 3 species. SABELLINAE. Radiles in spirals; eyes present, stylodes and webbing absent. Collar two- or four-lobed. Number of thoracic segments variable. Spatulate thoracic noto setae absent; late thoracic notopodia with iancent-shaped, transversely striated setae. Branchiomma Klliker 1858, Amphitrite bombyx Dalyell 1853; 21 species. SABELLINAE. Radiles sometimes spiralled, sometimes in semicircles; eyes and external stylodes present, webbing absent. Collar well developed. Thoracic spatulate setae absent; thoracic neuropodial companion setae absent. Eight thoracic segments Chone Kryer 1856, C. infundibuliformis Kryer 1856; 26 sf)ecies. FABRICINAE. More than five, usually many pairs of radioli present; distinct webbing present between the radioU. Collar well developed. Abdominal uncini with short, quadrangular base and crested tip. Abdomen with several setigers. Demonax Kinberg 1867b, D. leucaspis Kinberg 1867b; 8 species. SABELLINAE. Radioli in spiral; eyes present; webbing and external stylodes absent. Collar bilobed. Spatulate notosetae and neuropodial companion setae present in thorax. Both thoracic and abdominal uncini with long handles, but very sharply and distinctly bent and distally crested.
Desdemona Banse 1957, D. ornata Banse 1957; only species. FABRICINAE. Three pairs of radioli, radiolar backs rounded, webbing absent. Collar divided dorsally and unhed ventrally. Abdominal uncini with short, quadrangular bases and teeth in two rows along one side. Abdomen with four to 12 setigers. Dialychone Claparde 1870a, D. acstica Claparde 1870a; only species. FABRICINAE. Twelve or more pairs of radioli; webbing absent. Collar well developed on both sides. Abdominal uncini with short, quadrangular base and teeth in two or more rows along one side. Abdomen with several setigers. Distylidia Hartman l%\,Disrylia rugosa Moore 1904b;
2 species.
SABELLINAE. Radicles spiralled; eyes present, external stylodes and webbing absent. Collar bilobed. Both thoracic and abdominal uncini long-handled, but sharply bent. Anterior ventrum irregularly rugose. Euchone Malmgren 1866, Sabella analis Kryer 1856; 30 species. FABRICINAE. Numbers of radiles variable; webbing present. Collar usually well developed. Last abdominal setigers flanged laterally to form a deep spoon-shaped cavity. Abdominal uncini avicular. Abdomen with numerous setigers. Euchonella Fauchald 1972, E. magna Fauchald 1972; only species. FABRICINAE. Twelve pairs of radioli; webbing and eyes absent. Collar well developed, highest ventrally. Pygidium expanded to a large dorsal horse-shoe covering the last seven setigers. Abdominal uncini avicular. Abdomen with numerous setigers. Eudistylia Bush 1904a, Sabella vancouveri Kinberg 1867b; 3 species. SABELLINAE. Radiles spiralled; eyes present, webbing and external stylodes absent. Collar fourlobed. Spatulate thoracic notosetae present. Abdominal uncini avicular. Euratella Chamberlin 1919c, Laonome salmacidis Claparde 1870a; 2 species. SABELLINAE. Radioli in semi-circles; webbing and eyes present; external stylodes absent. Collar reduced.
1976
li9
Spamlate thoracic notosetae and neuropodial thoracic companion setae absent. Both thoracic and abdominal uncini avicular, with short, strongly bent handles. Fabricia Blainville 1828, Amphicora sabella Ehrenberg 1837; 18 species. FABRICINAE. Three pairs of radioli and branchial hearts present. Collar often very low dorsally and united ventrally. Abdomen with long-handled uncini. Three abdominal setigers. Fabrkiola Friedrich 1939, Manayunkia pacifica Annenkova 1934; 6 species. FABRICINAE. Three pairs of radiles and branchial hearts present. Collar distinct dorsally, united ventrally. Abdominal uncini short-handled. Three abdominal setigers. Fabrisabella Hartman 1969, F. vasculosa Hartman 1969; 2 species. FABRICINAE. Radiles in semi-circles; webbed, eyes and external stylodes absent. Collar high, but deeply and widely separated dorsally. Spatulate thoracic notosetae present; thoracic neuropodial companion setae absent. Thoracic uncini long-handled and gently curved; abdominal ones strongly bent. Numerous pairs of radiles present; abdomen with numerous setigers. Hypsicomus Grube 1870b, Sabella phaeotaenia Schmarda 1861; 12 species. SABELLINAE. Radicles in semi-circles; webbing, and external stylodes absent; eyes present. Collar well developed. Spatulate thoracic notosetae present; first notosetae in a long, straight or gently curved row. Abdominal uncini avicular. Jasmineira Langerhans 1880, J. caudata Langerhans 1880; 13 species. FABRICINAE. Eight or more pairs of radioli; webbing and eyes absent. Collar well developed. Abdominal uncini avicular; several abdominal setigers present. Laonome Malmgren 1866, L. kroeyeri Malmgren 1866; 4 species. SABELLINAE. Radiles in semi-circles; external stylodes and webbing absent; eyes usually present. Collar bilobed. Spatulate thoracic notosetae present; thoracic neuropodial companion setae absent. Both thoracic and abdominal uncini with short, flattened base and avicular head. Manayunkia Leidy 1859, M. speciosa Leidy 1859; 10 species. FABRICINAE. Two pairs of radioli, ventrally in addition a pair of palplike, vascularized smooth filaments. Webbing absent. Collar well developed; abdomen with three setigers; abdominal uncini long-handled, Megalomma Johansson 1926, Amphitrite vesiculosa Montagu 1815; only species.
SABELLINAE. Radiles in semi-circles; external stylodes and webbing absent. Very large, compound eyes present subdistally on a few dorsal radioli, otherwise absent. Collar two- or four-lobed. Spatulate thoracic notosetae present. Abdominal uncini avicular. Monroika Hartman \')5\h, Manayunkia africana Monro 1939b; only species. FABRICINAE. Six pairs of radiles, in part united by a web. Collar ventrally high, dorsally incised. Vascularized filaments absent. Two abdominal setigers; abdominal uncini long-handled. Myxicola Koch in Renier 1847, Terebella inndibulum Renier 1804; 5 species. MYXICOLINAE. Radiles in semi-circles, strongly webbed, external stylodes and eyes absent. Thoracic uncini minute and long-handled. Abdominal uncini avicular, in nearly complete cinctures around the posterior part of the body. Tube mucoid. Oriopsis Caullery and Mesnil 1896, Fabricia armandi Claparde 1864; 22 species. FABRICINAE. Three to five pairs of radiles; radiles externally flanged, webbing and eyes absent. Collar divided dorsally and fused ventrally. Abdomen with four or more setigers; abdominal uncini with short, quadrangular handles and teeth in several rows along one margin. Panousea Rullier and Amoureux 1970, P, africana Rullier and Amoureux 1970; only species. SABELLINAE. Radiles in semi-circles. Webbing, eyes and external stylodes absent. Collar four-lobed. Thoracic uncini long-shafted, neuropodial companion setae present. Potamethus Chamberlin 1919c, Potamilla malmgreni Hansen 1878; 10 species. SABELLINAE. Radiles in semi-circles; eyes, external stylodes and webbing absent. Collar very low, except ventrally where it is produced into a pair of triangular lobes. Thoracic uncini with long, nearly straight handles; abdominal uncini short-stemmed. Companion setae present. Potamilla Malmgren 1866, Sabella neglecta Sars 1850; 30 species, SABELLINAE. Radiles in semi-circles; external stylodes and webbing absent; eyes present. Collar two- or four-lobed. Spatulate thoracic notosetae present. Thoracic uncini short-handled, Potaspina Hartman 1969, P. pacifica Hartman 1969; only species. SABELLINAE. Radiles in semi-circles; eyes, external stylodes and webbing absent. Collar low dorsally, high and bifid ventrally. Anterior thoracic neuropodia with avicular uncini; last thoracic neuropodia with
140
Science Series 28
thick, acicular spines. Spatulate setae and companion setae present. Abdominal uncini small, acicular. Pseudobranchiomma Jones 1962, P. emersoni Jones 1962; only species. SABELLINAE. Radiles in semi-circles; eyes absent, webbing absent; external stylodes present, but small. Thoracic uncini short-handled, companion setae absent. Four thoracic setigers. Pseudofabricia Cantone 1972, P. aberrans Cantone 1972; only species. FABRICINAE. Eight radiles on a greatly prolonged, distally bifid anterior end and an extended rounded, lower lip. Abdominal uncini with short base and several rows of teeth. Three abdominal setigers. Sabella Linnaeus 1767, 5. penicillus Linnaeus 1767; 35 species. SABELLINAE. Radiles in semi-circles; eyes present, external stylodes and webbing absent. Collar four-lobed. Spatulate thoracic notosetae absent. Sabellastarte Savigny 1818, Eurato sanctijosephi Gravier 1906; 8 species. SABELLINAE. Radiles spiralled; eyes present, webbing and external stylodes absent. Collar well developed, widely separated dorsally. Spatulate and companion setae absent. Schizobranchia Bush 1904a, S. insignis Bush 1904a; 2 species. SABELLINAE. Radicles in semi-circles; dichotomously divided. Eyes present; external stylodes and webbing absent. Collar four-lobed. Thoracic uncini long-handled, but bent; companion setae present. Spirographis Viviani 1805, S. spallanzani Viviani 1805, 4 species. SABELLINAE. Radiles spiralled, with one half very much larger than the other, only one part spiralled. Stylodes and webbing absent; eyes present. Collar four-lobed. All uncini avicular. Taxonomic Notes The sub-families as accepted here are based on the structure of the neuropodial uncini of the thorax, in that members of the FABRICINAE have gently curved, long-handled uncini, and members of SABELLINAE have strongly bent, avicular uncini. However, all sabellids with thoracic neuropodial companion setae have been included in the SABELLINAE, irrespective of the structure of the uncini. The genus Pseudopotamilla Bush is considered here a junior synonym of Potamilla and the genus Trichosobranchella Dybowski 1929, is referred to Manayunkiu. Jones (1974a) revised the original material of Monroika and pointed out that the character used to separate
this genus from all other FABRICIIN genera is incorrect; it does not have the webbed tentacular crown claimed by Hartman. Members of this genus will key out with Manayunkia above. It differs from the latter in that it has short limbate setae in both the first setigers; Manayunkia has such setae only in the first setiger and they are much longer. Furthermore, Monroika has two rather than three abdominal setigers. Jones (1974a) also described a new genus, Brandtika (genotype: B. asitica), which will key out in the same complex. It differs from other genera in this complex in that it has pilose setae in the last three thoracic setigers. This genus could not be included in the key, since it was based on dried material, and no information was available about the structure of the tentacular crown. Invalid Genera Amphicora Ehrenberg 1837, see Fabricia Amphicorina Quatrefages 1866, see Oriopsis Anamobaea Kryer 1856, see Hypsicomus Arippasa Johnston 1865, see Myxicola Aspeira Bush 1904a, see Potamilla Branchiomma Claparde 1870a, seeMegalomma Chaponella Rullier 1972, see Euchone Dasycharte Sars 1862, see Branchiomma Dasychonopsis Bush 1904a, see Branchiomma Distylia Quatrefages 1866, seeBispira Dybowscella Nusbaum 1901, see Manayunkia Eriographis Grube 1850, see Myxicola Eurato Saint-Joseph 1894, see Hypsicomus Fabriciella Zenkevitch 1935, see Fabriciola Garjaiowella Dybowskii 1929, see Manayunkia Gorbunovia Annenkova 1952, see Potamethus Gymnosoma Quatrefages 1866, see Myxicola Haplobranchus Bourne 1883, see Manayunkia Hypsicomatides Augener 1922, see Hypsicomus Hypsicomatopsis Augener 1922, see Hypsicomus Laonomedes Chamberlin 1919c, see Potamilla Leiobranchus Quatrefages 1850, see Myxicola Leptochone Claparde 1870b, see Myxicola Megachone Johnson 1901, see Chone Metachone Bush 1904a, see Chone Metalaonome Bush 1904a, seeBispira Notaulax Tauber 1879, see Hypsicomus Oria Quatrefages 1866, see Oriopsis Oriades Chamberlin 1919c, see Oriopsis Oridia Rioja 1917, see Oriopsis Othonia Johnston 1835, questionably Fabricia Parachonia Kinberg 1867b, see Chone Parasabella Bush 1904a, see Demonax Potamis Ehlers 1887, see Potamethus Protulides Webster 1884, see Hypsicomus Pseudopotamilla Bush i904a, see Potamilla Sabellina Dujardin 1839a, indeterminable Sabina Williams 1851, indeterminable
976
141
Sitophaga Gistel 1848, see Fabricia Trichosobranchella Dybowski 1929, see Manayunkia
FAMILY CAOBANGIIDAE JONES
1974b
Small, short-bodied sabelform polychaetes with three pairs of radioli in a tentacular crown. Digestive tract U-shaped with the anus opening dorsally and far anteriorly on the body. First setiger with neurosetal palmate hooks, remainder of the thoracic setigers without hooks; two kinds of avicular hooks present in a posterior region. This fainily was recently proposed by Jones (1974b) for a series of small polychaetes that live in close association with molluscs in freshwater in Southeast Asia. Most of the forms burrow in the shell of the host. The family consists of a single genus, Caobangia Giard 1893, with type-species C. billeti Giard 1893 and a total of seven known species. Up to this time, the genus has been considered among sabellid enigmaticae, but the demonstration of a series of forms spread over a larger geographical region, made the recognition of a new family necessary. 1969
The family is known for a single genus and species, Sabellonga disjuncta Hartman 1969 from northern Baja California. The specimen resembles a sabellid that has lost its tentacular crown. However, there is no trace of the loss of a tentacular crown, or that such has ever been present. The presence of the giant falcate spines in far posterior setigers also is characteristic. Otherwise, the setal equipment is largely what one would expect in a member of the SABELLINAE.
FAMILY SERPULIDAE JOHNSTON
1865
FAMILY SABELLONGIDAE HARTMAN
Body cylindrical with few thoracic and many abdominal segments. Peristomium forms a bevelled collar around the prostomium (tentacular crown absent). Setae include long-handled uncini with companion setae and giant falcate spines.
Body separated into two regions; a thorax with a thoracic membrane (absent in rare instances) and dorsal capillary or limbate setae and an abdomen with ventral setae and dorsal uncini. One radile often transformed into an operculum. Tube calcareous. The serpulids and the closely allied spirorbids represent a sub-specialty of their own within the polychaetes. The family has been reviewed on a couple of occasions; Saint-Joseph (1894) published an extensive review and Southward (1963) gave a key to all known genera. The generic sub-divisions within the family are nevertheless debatable. The older taxonomic groupings placed emphasis on the structure of the operculum and on the overall body-construction (development of the thoracic membrane and collar, presence or absence of setae in specified segments, etc.). More recently, some authors (especially Zibrowius) have placed more emphasis on the detailed structure of the uncini and setae. The key below represents a compromise, and may, as such, be difficult to use.
Key to Genera la. lb. 2a (la). 2b (la). 3a (2b). 3b (2b). 4a (3a). 4b (3a). 5a (4b). 5b (4b). 6a (3b). 6b (3b). 7a (lb). 7b (lb). 8a (7a). 8b (7a). 9a (8a). 9b (8a). 10a (8b). Operculum absent 2 Operculum present 7 Abdominal setae trumpet-shaped Pseudoserpula Abdominal setae marginally dentate and geniculate, limbate or capillary 3 Slender capillary or limbate collar setae present 4 Collar setae modified 6 Thoracic membrane absent Salmacinopsis Thoracic membrane present 5 Thoracic membrane reaches setiger 5 Subprotula Thoracic membrane reaches setiger 7 Salmacina Abdominal setae geniculate Salmacina Abdominal setae slender, nearly straight capillaries Protis Operculum carried on a branchial radile 8 Operculum carried on a modified stalk 12 Five thoracic setigers present 9 Six or more thoracic setigers present 10 Collar setae slender and limbate; long asetose region present between thorax and abdomen Josephella Collar setae basally dentate; thorax and abdomen not separated by a long asetose region Dipomatus Collar setae simple and tapered limbate Apomatus
142 lOb (8b). I la (10b). 1 lb (lOb). 12a (7b). I2b (7b). 13a ( 12b). 13b (I2b). 14a (13a). 14b (13a). 15a (14a). 15b (14a). 16a (I5b). 16b (15b). 17a (13a). 17b (13a). 18a (17a). 18b (17a). 19a (17b). 19b (17b). 20a (19a). 20b (19a). 21a (20b). 21b (20b). 22a (21b). 22b (21b). 23a (19b). 23b ( 19b). 24a (23a). 24b (23a). 25a (23b). 25b (23b). 26a (13b). 26b (13b). 27a (26a). 27b (26a). 28a (27b). 28b (27b). 29a (28a). 29b (28a). 30a (28b). 30b (28b). 31a (30b). 31b (30b). 32a (31b). 32b (31b). 33a (26b). 33b (26b). 34a (33b). 34b (33b). 35a (34b). 35b (34b). 36a (35a).
Science Series 28
Collar setae with a few coarse teeth basally and a Hmbate denticulate blade distally II Operculum a shallowly depressed cone on a spherical swelling of the opercular stalk ... Filogranula Operculum a depressed cone; stalk without the subdistal swelling Filograna Opercular stalk flattened and ribbonlike Mesavermilia Opercular stalk oval or circular in cross-section (sometimes with wings or spines) 13 Opercular stalk with wings or spines 14 Opercular stalk without wings or spines 26 Opercular stalk with spines 15 Opercular stalk with wings (sometimes produced into points distally) 17 Opercular stalk with spines on one side only; six thoracic setigers present Spirodiscus Opercular stalk with four spines in a cross; seven thoracic setigers present 16 Thoracic neuropodia widely separated anteriorly and approaching posteriorly; collar setae all of one kind, with basal pilose boss and distal spines Spirobranchus Thoracic neuropodia equidistant in all setigers; collar setae of two kinds; either capillary or with a smooth double-boss basally and smooth tapering tips Crucgera Collar setae absent 18 Collar setae present 19 Operculum with two projections Olga Operculum without projections Pomatoleios Collar setae limbate or capillary 20 Collar setae with two separate limbations (bayonet-type), with a basal boss or setose 23 Anterior abdominal setae stout, acute spines Paumotella Anterior abdominal setae trumpet-shaped or geniculate 21 Opercular cap black and chitinous Crosslandiella Opercular cap calcareous 22 Opercular cap flat, with or without spines Pomatoceros Operculum distally excavated, bordered by two eccentrically placed thorns connected by a low ridge Pseudopomatoceros At least anterior abdominal setae trumpet-shaped 24 Anterior abdominal setae geniculate 25 Operculum conical, collar setae basally pilose Conopomatus Operculum with slanting distal plate, collar setae bayonet-shaped Temporaria Uncini with anterior peg pointed Omphalopomopsis Uncini with anterior peg gouge-shaped (hollowed out from beneath) Pomatostegus Collar setae absent 27 Collar setae present 33 Abdominal setae absent; abdominal uncini present Rhodopsis Abdominal setae present; uncini usually present 28 At least some abdominal setae geniculate 29 All abdominal setae slender capillaries 30 Uncini with numerous teeth Placostegus Uncini with seven to nine teeth Marifugia Abdominal uncini absent Bonhourella Abdominal uncini present 31 Tube free, tusk-shaped and smooth Ditrupa Tube at least partly attached 32 Opercular stalk calcified; operculum funnel-shaped Scierostyla (in part) Opercular stalk not calcified; operculum spherical Dasynema Opercular spines movable Galeolaria Opercular spines, if present, immovable 34 Abdominal setae slender capillaries Schiiocraspedon Abdominal setae trumpet-shaped or geniculate 35 Abdominal setae trumpet-shaped 36 Abdominal setae geniculate 39 Operculum a simple funnel 37
/97(5 36b (35a). 37a (36a). 37b (36a). 38a (36b). 38b (36b). 39a (35b). 39b (35b). 40a (39a). 40b (39a). 41a (40a). 41b (40a). 42a (41a). 42b (41a). 43a (41b). 43b (41 b). 44a (43b). 44b (43b). 45a (40b). 45b (40b). 46a (45a). 46b (45a). 47a (46b). 47b (46b). 48a (45b). 48b (45b). 49a (39b). 49b (39b). 50a (49b). 50b (49b). 51a (50a). 51b (50a). 52a 52b 53a 53b 54a 54b (51a). (51a). (52b). (52b). (53b). (53b).
43
Operculum with a basal funnel and in addition various spines or hoods forming a distal part .... 38 Collar setae basalty minutely hirsute Paraserpula Collar setae basally dentate Serpula Operculum with a large, glandular hood-shaped distal part Olgaharmania Operculum with series of distal spines and, sometimes, smaller hoods Hydroides Side setae {Apomatus-ssX?is.) absent 40 Side setae present 49 Collar setae with few coarse teeth in one or two marginal rows 41 Collar setae at least in part limbate 45 Operculum distally ornamented 42 Operculum distally smooth, rounded or excavate 43 Operculum with concentric series of teeth, nearly radial in structure Neopomatus Operculum with a single series of long, strong teeth, making it bilaterally symmetrical . Mercierella Collar setae with teeth in two rows Sphaeropomatus Collar setae with teeth in a single row 44 Operculum distally rounded Ficopomatus Operculum distally excavate Mercierellopsis At least some collar setae with a basal dentate or hirsute region 46 All collar setae limbate 48 Simple limbate collar setae present, in addition to some with a basal group of spines Omphalopoma All collar setae with basal denticulated or hirsute region 47 Six thoracic setigers present Hyalopomatus Seven thoracic setigers present Pseudochitinopoma Opercular stalk calcified; abdomjnal setae with only a short geniculate tip Sclerostyla (in part) Opercular stalk not calcified; abdominal setae with about one-half of the exposed setae in the geniculate tip Neovermilia Opercular stalk annulated Calcareopomatus Opercular stalk not annulated 50 Collar setae limbate 51 Collar setae with a basal boss in addition to the limbate distal portion 52 Both thoracic and abdominal uncini with all teeth in a single row; anterior peg entire .. Vermiliopsis Thoracic uncini with teeth in a single row, anterior peg furcate; abdominal uncini with teeth in several rows Pseudovermilia Both thoracic and abdominal uncini with teeth in several rows Filogranula (in part) Thoracic uncini with teeth in a single row 53 Operculum distally rounded Chitinopomoides Operculum distally excavate 54 Opercular stalk with three distal bulbs Janita Opercular stalk without swellings Chitinopoma
Generic Dfinitions Apomaius Philippi 1844, A, ampulliferus Philippi 1844; 8 species. SERPULINAE. Seven thoracic setigers; globular operculum on a radile. Collar setae simple limbates; side setae present. Uncini with teeth in several rows in both thorax and abdomen; anterior peg very long. Bonhourella Gravier 1905b, B. insignis Gravier 1905b; only species. SERPULINAE. Six thoracic setigers; operculum flat or lightly convex, opercular stalk distinct. Collar setae absent; abdominal setae capillary; abdominal uncini
absent. Thoracic uncini with anterior peg gougeshaped. Calcareopomatus Straughan 1967a, C. dewae Straughan 1976a; only species. SERPULINAE. Seven thoracic setigers; operculum with a globular base and a flat, calcareous plate; stalk annulated. Collar setae small and limbate; side setae present. Thoracic uncini with anterior peg bifid. Chitinopoma Levinsen 1884, Hydroides norvegica groenlandica Mrch 1863; 2 species. SERPULINAE. Seven thoracic setigers; operculum conical without wings or spines on the stalk. Collar
144
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setae of fin and blade construction; side setae present; abdominal setae geniculate with triangular blade. Thoracic uncini with teeth in a single row; abdominal ones with teeth in several rows. Chitinopomoides Benham 1927, C. wilsoni Benham 1927; only species. SERPULINAE. Seven thoracic setigers; operculum rounded, stalk without spines or wings. Collar setae with large boss below a limbate zone; side setae present; abdominal setae geniculate with long triangular blade. Thoracic uncini with teeth in a single row; abdominal ones with teeth in several rows. Conopomatus Pillari 1960, C. acuiconus Pillai 1960; 2 species. SERPULINAE. Seven thoracic setigers; operculum conical, stalk winged. Collar setae present, pilose at base; side setae present; other thoracic setae limbate; abdominal setae trumpet-shaped. Crosslandiella Monro 1933, C. multispinosa Monro 1933; 2 species. SERPULINAE. Seven thoracic setigers; operculum with black, chitinous plate, surmounted by a spinose column; stalk winged. Collar setae lancet-shaped and tapering; side setae present. Abdominal setae geniculate. Crucgera Benedict 1887, C. websteri Benedict 1887; 5 species. SERPULINAE. Seven thoracic setigers; operculum with simple funnel, distal end of stalk with four large spines forming a cross. Collar setae limbate; side setae absent; abdominal setae trumpet-shaped. Uncini with few teeth; anterior peg very large. Dasynema Saint-Joseph 1894, Serpula chrysogyrus Grube 1878; only species. SERPULINAE. Seven thoracic setigers; operculum spherical, opercular stalk smooth. Collar setae and side setae absent; all thoracic setae limbate, abdominal ones capillaries. Dipomatus Ehlers 1913, D. serpulides Ehlers 1913; only species. SERPULINAE. Five thoracic setigers; two operculae with branchial filaments on peduncle; each distally funnel-shaped and marginally dentate. Collar-setae basally dentate. Ditrupa Berkeley 1835, Dentalium arietinum O.F. Mller 1776; 8 species. SERPULINAE. Tube free, tusk-shaped; operculum an inverted cone with a chitinous plate. Collar setae absent; thoracic setae limbate and capillaries; abdominal setae capillaries. Uncini with numerous teeth in several rows. Ficopomatus Southern 1921, F. macrodon Southern I92I; 2 species.
FICOPOMATINAE. Seven thoracic setigers; operculum pear-shaped, soft or chitinous; stalk smooth. Some collar setae with dentate boss, others serrated limbates. Side setae absent. Abdominal setae geniculate with a dentate, slender tip. Uncini with few teeth in a single row, Filograna Oken 1815, F. implexa Berkeley 1828; only species. FILOGRANINAE. Six to 12 thoracic setigers; rounded operculum on one of the radiles. Collar setae notched with limbate expansion at the base. Side setae and limbate setae present in thorax; abdomen with geniculate setae. Uncini with numerous teeth in several rows. Filogranula Langerhans 1884, F. gracilis Langerhans 1884; 3 species. FILOGRANINAE. Seven thoracic setigers; operculum a small cone on a small, spherical base; stalk either frondose or smooth; only four radicles present. Collar setae present; side setae present; uncini with numerous teeth in several rows. Galeolaria Savigny 1818, G. caespitosa Savigny 1818; 5 species. SERPULINAE. Seven thoracic setigers; operculum with calcareous plate surmounted by movable spines; opercular stalk winged. Collar setae present and very short and slender. Hyalopomatus Marenzeller 1878, H. claparedii Marenzeller 1878; 4 species. SERPULINAE. Six thoracic setigers; operculum bladder-shaped; stalk smooth. Collar setae of fin and blade construction; side setae absent. Abdominal setae with only the distalmost tip geniculate. Uncini with teeth in several rows; anterior peg furcate. Hydroides Gunnerus 1768, H. norvegica Gunnerus 1768; 85 species. SERPULINAE. Seven thoracic setigers; operculum basally with a marginally dentate funnel, distally with a crown of spines or smaller hoods, or a second, distal funnel present; stalk without spines or wings. Collar setae limbate; side setae absent; abdominal setae trumpet-shaped. Janita Saint-Joseph 1894, Serpula fimbriata delle Chiaje 1828; only species. SERPULINAE. Seven thoracic setigers; operculum infundibular with three large distal bulbs on the stalk; opercular plate chitinous with a single large spine. Collar setae basally dentate; side setae present; abdominal setae geniculate. Josephella Caullery and Mesnil 1896, J. marenzelleri Caullery and Mesnil 1896; 2 species. SERPULINAE. Five thoracic setigers; operculum at end of normal radile, rounded. Collar setae slender
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limbate; uncini with double rows of teeth; anterior peg deeply bifid. Marifugia Absalon and Hrabe 1930, M. cavatica Absalon and Hrabe 1930; only species. SERPULINAE. Six thoracic setigers; operculuni conical and distally smooth; stalk without spines or wings. Collar setae and side setae absent; thoracic setae slender and straight; abdominal setae geniculate. Uncini with teeth in a single row, maximally nine teeth present in thoracic uncini. Membranopsis Bush 1910, A. inconspicua Bush 1910; only species. SERPULINAE. Nine thoracic setigers; operculum unknown. Thoracic membrane fused over the back at ninth seliger; collar setae limbate. Uncini similar to those m Protula. Mercierelia Fauvel 1923b, M. enigmtica Fauvel 1923b; only species. FICOPOMATINAE. Seven thoracic setigers; operculum distally oblique with chitinous plate covered by series of curved spines. Collar setae marginally dentate; side setae absent; uncini with teeth in one row in thorax, partially in two rows in abdomen. Merciereliopsis Rioja 1945, M. prietoi Rioja 1945; only species. FICOPOMATINAE. Seven thoracic setigers; operculum smooth, distally cup-shaped; stalk smooth. Collar setae dentate; side setae absent. Uncini with few teeth (7-12), anterior peg undercut. Metavermilia Bush 1904a, Vermilia multicristata Philippi 1844; 4 species. SERPULINAE. Seven thoracic setigers; operculum bladder-shaped; stalk flattened, ribbon-shaped. Collar setae limbate; side setae present; abdominal setae genicdate or capillaries. Thoracic uncini with teeth in a single row; abdominal ones with teeth in one or several rows. Neopomatus Pillai 1960, N. uschakovi Pillai 1960; 2 species. FICOPOMATINAE. Seven thoracic setigers; operculum with series of concentric rows of teeth, nearly radial in structure; stalk smooth. Collar setae dentate; side setae absent. Uncini with few teeth. Neovermilia Day 1961, N. capensis Day 1961; 2 species. SERPULINAE. Seven thoracic setigers; operculum spherical or slightly cup-shaped; stalk smooth. Collar setae limbate; side setae absent. Abdominal setae geniculate, finely dentate capillaries. Thoracic uncini with few teeth in a single row. Olga Jones 1962, O. elegantissima Jones 1962; only species.
SERPULINAE. Six thoracic setigers; operculum with a central tooth and paired lateral horns, stalk winged. Collar setae absent, side setae absent; thoracic setae limbate; uncini with about twenty teeth; anterior peg gouge-shaped. Olgaharmania Rioja 1941b, Hydroides glandiferum Rioja 1941a; only species. SERPULINAE. Seven thoracic setigers; operculum with basal corona of spines surmounted by very large, glandular chitinous hood equipped on one side with four large spines. Collar setae basally with paired bosses; side setae absent. Uncini with numerous teeth; anterior peg very large. Omphalopoma Morch 1863, O. umbilicata Mrch 1863; 2 species. SERPULINAE. Six or seven thoracic setigers; operculum cup-shaped, sometimes ornamented; stalk smooth. Collar setae of two kinds, simple limbates and fin and blade setae. Side setae present. Thoracic uncini with teeth in a single row; abdominal ones with teeth in several rows; anterior peg furcate. Omphalopomopsis Saint-Joseph 1894, Omphalopoma langerhansii Marenzeller 1884; 2 species. SERPULINAE. Seven thoracic setigers. Operculum distally smooth; opercular stalk with wings. Collar setae of two kinds, slender capillaries and limbate with a pilose subdistal boss. Side setae present; abdominal setae geniculate with a triangular blade. Thoracic uncini with teeth in a single row; anterior peg entire. Paraserpula Southward 1963, P. planorbis Southward 1963; only species. SERPULINAE. Seven thoracic setigers; operculum funnel-shaped, marginally dentate; stalk smooth. Collar setae bayonet-shaped and finely pilose; side setae absent; abdominal setae trumpet-shaped. Thoracic uncini with teeth in a single row; abdominal ones with teeth in several rows; anterior peg large. Paumotella Chamberlin 1919c, P. takemoana Chamberlin 1919c; only species. SERPULINAE. Seven thoracic setigers present; operculum conical and distally flattened, margin entire, stalk with wings. Collar setae limbate; side setae absent; anterior abdominal setae stout acute spines; posterior ones fine capillaries. Placostegus Philippi 1844, Serputa tridentala Fabricius 1780; 18 species. SERPULINAE. Seven thoracic setigers; of)erculum funnel-shaped with chitinous distal plate; stalk smooth. Collar setae absent; abdominal setae geniculate with widely triangular dentate distal plates. Uncini with numerous teeth.
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Pomatoceros Philippi 1844, Serpula triquetra Linnaeus 1767; 13 species. SERPULINAE. Seven thoracic se tigers; operculum a flat calcareous plate, with or without spines; stalk with broad wings. Collar setae small and limbate; thoracic setae limbate; abdominal setae trumpet-shaped. Side setae absent. Uncini with numerous teeth. Pomatoleios Pixell i913b, P. crosslandi Pixell 1913b; 3 species. SERPULINAE. Seven thoracic setigers; operculum distally flat; stalk winged. Collar setae and side setae absent; abdominal setae trumpet-shaped with one long spine at one side. Uncini with about ten teeth in both thorax and abdomen. Pomatostegus Schmarda 1861, P. macrosoma Schmarda 1861; 8 species. SERPULINAE. Seven thoracic setigers; operculum with chitinous plate surmounted by a column with several discs; stalk winged. Collar setae with subdistal pilose boss; side setae present; abdominal setae geniculate. Uncini with teeth in a single row; anterior peg simple. Protis Ehlers 1887, P. simplex Ehlers 1887; 3 species. SERPULINAE. Seven thoracic setigers; operculum absent. Collar setae of fin and blade construction with basal dentate fm well separated from the blade. Side setae present; abdominal setae nearly capillary. Thoracic uncini with teeth in a single row; abdominal ones with teeth in several rows. Protula Risso 1826, Serpula tubularia Montagu 1803 in Mclntosh 1923; 23 species. SERPULINAE. Seven thoracic setigers; operculum absent. Collar setae simple limbates; side setae present. Uncini with teeth in several rows; anterior peg very long. Pseudochitinopoma Zibrowius 1969, Hyalopomatopsis occidentatis Bush 1904a; only species. SERPULINAE. Seven thoracic setigers; operculum bladder-shaped; stalk without wings. Collar setae with fin and blade construction and little separation between the basal fin and the distal blade. Side setae absent; abdominal setae geniculate with narrow blade. Thoracic uncini with teeth in a single row; anterior peg furcate; abdominal uncini with teeth in several rows, Pseudopomatoceros Holly 1936, Pomatoceropsis roxasi Holly 1935; only species. SERPULINAE. Seven thoracic setigers; operculum distally cup-shaped with two eccentrically placed thorns connected by a low ridge; stalk winged. Collar setae fine capillaries; side setae absent, thoracic setae limbate, abdominal setae with flat crown and a long spike. Uncini with numerous teeth; anterior peg furcate.
Pseudoserpula Straughan 1967b, P. rugosa Straughan 1967b; 2 species. SERPULINAE. Seven thoracic setigers; operculuin absent. Collar setae limbate; side setae absent; abdominal setae trumpet-shaped. Uncini with few teeth; anterior peg large, blunt. Pseudovermilia Bush 1907b, Spirohranchus occidenlalis Mclntosh 1885; 5 species. SERPULINAE. Seven thoracic setigers; operculum distally ornamented; stalk smooth. Collar setae limbate, side setae present; abdominal setae geniculate with triangular blades. Thoracic uncini with teeth in a single row; anterior peg furcate; abdominal ones with teeth in several rows. Rhodopsis Bush 1904a, R. pusillus Bush 1904a; only species. SERPULINAE. Seven thoracic setigers; operculum covered distally with chitinous plate with spines arranged in a rosette; shaft smooth, Collar setae absent; abdominal setae absent, Uncini with teeth in a single row, Salrrmcina Claparde 1870a, S. incrustans Claparde 1870a; 8 species. FILOGRANINAE. Five to nine thoracic setigers; operculum absent. Collar setae notched with limbate subdistal and distal parts; side setae present. Abdominal setae geniculate. Uncini with teeth in several rows. Salmacinopsis Bush 1910, S. setosa Bush 1910; only species. FILOGRANINAE. Nine thoracic setigers; operculum absent. Collar setae capillary; other thoracic setae capillary and limbate with a few serrate setae in posterior thoracic setigers; abdominal setae geniculate and serrated. Uncini with numerous very fine teeth; anterior peg large and blunt. Thoracic membrane absent, Schizocraspedon Bush 1904a, Serpula furcifera Grube 1878; only species. SERPULINAE. Seven thoracic setigers; operculum on a modified radile; distally with a chitinous cover consisting of two concentric funnels, one inside the other; both marginally split with long, slender processes. Collar setae present. Abdominal setae capillary; superior thoracic setae geniculate with conspicuous spines at the base. Sclerostyla Mrch 1863, 5, ctenaclis Mrch 1863; 2 species. SERPULINAE. Seven thoracic setigers; operculum funnel-shaped; opercular stalk smooth, calcified, collar setae present or absent, if present finely limbate; side setae absent; abdominal setae nearly capillary, with very short geniculate tips. Thoracic uncini with teeth
\97f>
THE
Poi.YCHAETE WORMS
147 Taxonomic Notes
in a single row; abdominal ones with teeth in several rows. Serpula Linnaeus 1767, S. vermicularis Linnaeus 1767; 17 species. SERPULINAE. Seven thoracic setigers; operculum funnel-shaped with crenulated margin; opercular stalk smooth. Collar setae either limbate or with a dentate boss; side setae absent. Abdominal setae trumpetshaped. Uncini with few teeth; anterior peg large. Sphaeropomatus Tread well 1934, S. miamiensis Treadwell 1934; only species. FICOPOMATINAE. Seven thoracic setigers; operculum soft and vesicular; stalk smooth. Collar setae spinose with teeth in two rows; side setae absent; other setae slender capillaries. Spirobranchus Blainville \&IS, Serpula gigantea Pallas 1766; 30 species. SERPULINAE. Seven thoracic setigers; operculum with calcareous distal plate, usually ornamented; opercular stalk winged. Collar setae bayonet-shaped and pilose; some forms have limbate collar setae; side setae absent. Abdominal setae trumpet-shaped. Thoracic uncini with teeth in a single row; anterior peg furcate. Thoracic neuropodia are separated widely anteriorly and approaching in late thoracic setigers. Spirodiscus Fauvel 1909, S. grimaidii Fauvel 1909; only species. FILOGRANINAE. Six thoracic setigers; operculum funnel-shaped on a small ampulla; stalk with dorsal spines. Collar setae flattened and lancet-shaped. Uncini with numerous teeth. Subprotula Bush 1910, S. longiseta Bush 1910; 2 species. SERPULINAE. Seven thoracic setigers; operculum absent. Collar setae capillaries. Uncini with about 15 teeth; anterior peg very large. Thoracic membrane to fifth setiger only; collar tri-lobed. Temporaria Straughan 1967b, Vermilia poiytrema Philippi 1844; 2 species. SERPULINAE. Seven thoracic setigers; operculum with slanting calcareous plate; stalk winged. Collar setae bayonet-shaped with both regions limbate, separated by a notch. Abdominal setae trumpet-shaped. Thoracic uncini with at least ten teeth. Vermiliopsis Saint-Joseph 1894, Serpula irifundibulum Linnaeus 1758; 35 species. SERPULINAE. Seven thoracic setigers; operculum rounded or flattened, stalk without wings. Collar and side setae present. Thoracic uncini with teeth in a single row; anterior peg entire; abdominal uncini with teeth in a single row.
The genus Membranopsis Bush 1910, included in the generic definitions, has been omitted from the key, since the structure of the operculum remains unknown. The genus Hydroides has been defined here to include Eupomatus Philippi 1844, as is customarily done by most specialists today. It may be of some value to recognize the difference between Hydroides SENSU STRICTU-dnd Eupomatus at the sub-generic level.
Invalid Genera Apomatopsis Saint-Joseph 1894, s^t Apomatus Cystopomatus Gravier 1911a, see Hyalopomatus Eucarphus Mrch 1863, see Hydroides Eupomatus Philippi 1844, see Hydroides Filigrana Mrch 1863, see Filograna Filopora Fleming 1825, e Filograna Glossopsis Bush 1904a, see Hydroides Helena Castelnau 1842, see Serpula Hyalopomatopsis Saint-Joseph 1894, see Hyalopomatus Microserpula Dons 1931, see Chitinopoma Paravermilia Bush 1907b, see Vermiliopsis Piratesa Templeton 1835, indeterminable Placostegopsis Saint-Joseph 1894, see Placostegus Podioceros Quatrefages 1866, see Pomatoceros Polyphragma Quatrefages 1866, see Hydroides Pomatoceropsis Gravier 1905b, see Spirobranchus Pomatoceropsis Holly 1935, see Pseudopomatoceros Proplacostegus Bush 1904a, see Placostegus Psymobranchus Philippi 1844, see Protula Siliquaria Savigny 1818, indeterminable Spiramella Blainville in Quatrefages 1866, see Protula Vermilia Savigny 1818, see Spirobranchus Zopyrus Kinberg 1867b, see Serpula 1970
FAMILY SPIRORBIDAE PILLAI
Body separated into two regions; a thorax with a thoracic membrane, dorsal setae and ventral uncini and an abdomen with ventral setae and dorsal uncini. One radile transformed into an operculum; tube calcareous. Body asymmetrical and tube coiled in a spiral; thorax with setigerous rudiments of three, four or five segments. The spirorbids are rather difficult to work with since identification requires precise and accurate work with compound microscopes; prior to that of course, the specimens must be dissected out from their tubes as completely as possible. Recent reviews by BaileyBrock, Knight-Jones and Vine has clarified the generic sub-division of the group considerably and is largely followed here (references to articles can be found be-
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low). Where the above-mentioned authors have yet to treat a genus. Pillai (1970) has been followed.
The family is most frequently considered a subfamily under the serpulids.
Key to Genera la. lb. 2a (la). 2b (la). 3a (2a). 3b (2a). 4a (3a). 4b (3a). 5a (lb). 5b (lb). 6a (5a). 6b (5a). 7a (6b). 7b (6b). 8a (7b). 8b (7b). 9a (8b). 9b (8b). 10a (5b). 10b (5b). 1 la (10a). With three or more pairs of thoracic tori 2 With two pairs of thoracic tori 5 With four or more pairs of thoracic tori or at least rudiments of a fourth pair present on the concave side 3 With three pairs of thoracic tori 17 With at least four complete thoracic tori present 4 With remnants of a fourth thoracic torus present 21 Four pairs of thoracic tori present Anomalorbis Approximately seven pairs of thoracic tori present Neomicrorbis Side setae absent 6 Side setae present 10 Margins of collar fused to form dorsal tunnel Dexiospira Margins of collar not fused 7 Dextrally coiled Pillaiospira Sinistrally coiled 8 Collar setae limbate Leodora Collar setae of fin and blade construction 9 Thoracic uncini with simple, blunt anterior peg Simplicaria Thoracic uncini with broadly indented anterior peg Eulaeospira Dextrally coiled 11 Sinistrally coiled 12 Collar setae of fin and blade construction Spirorbella
gf**^
38. (A), Family SPIRORBIDAE, Laeospira sp., simplified after Day 1964, 15x; (B), Protolaeospira patagnica, tube, after Day 1967, about 15x; (C), Family SERPULDAE, Spirobranchus spinosus, after Hartman, 1969, 5x; (D), Serpula vermicularis, tube, natural size.
FIGURE
97(, lib (lOa). 12a (10b). 12b ( 10b). 13a (12a), 13b (12a). 14a (13b). 14b (13b). 15a (12b). 15b (12b). 16a (15b). 16b (15b). 17a (2b). 17b (2b). 18a (17a). 18b (17a). 19a (18b). 19b (18b). 20a (17b). 20b (17b). 21a (3b). 21b 22a 22b 23a 23b 24a 24b 25a 25 b (3b). (21b). (21b). (22a). (22a), (22b), (22b). (24b). (24b),
149
Collar setae simply limbate Janua Collar setae simply limbate 13 Collar setae of fin and blade construction 15 Collar fused dorsally to form tunnel Romanchella Collar not fused dorsally 14 Incubation of larvae in opercutar chamber; abdominal setae with blades as those of the collar setae Fauveldora Incubation in the tube; abdominal setae very small with short blades Buskiella Thoracic uncini with several rows of teeth Spirorbis Thoracic uncini with a single row of teeth 16 Two or more opercular plates on top of each other on the opere ulum Duplicara A single opercular plate present Pileolaria Side setae present 18 Side setae absent 20 Collar setae of fin and blade construction Paralaeospira Collar setae simply limbate 19 Incubation of larvae in opercular chamber Protoleodora Larvae attached to stalk in faecal groove Metalaeospira Tube tightly coiled along its whole length Circeis Tube coiled only basally, most of the tube uncoiled Helicosiphon Three thoracic tori and four fascicles of setae on the convex side; four thoracic tori and five fascicles of setae on the concave side Ampiara Fourth setiger represented by a rudiment of the torus on the concave side 22 Sinistrally coiled 23 Dextrally coiled 24 Side setae present Protolaeospira Side setae absent Capeospira Collar setae simply limbate Prodexiospira Collar setae of fin and blade construction 25 Thoracic uncini with furcate anterior peg Dextralia Thoracic uncini with bluntly rounded anterior peg Paradexiospira generic level, as indicated in Table 3. I believe that this difference in approach is relatively unimportant for the time being.
The key given above suggests that all named taxa are at the same level. Vine (1972) and Knight-Jones (1973) used several of the contained taxa at the sub-
TABLE
Alphabetic list of genera and sub-genera of Spirorbidae as suggested by Vine (1972) and Knight-Jones (1973). Genus Ampiara Anomalorbis Capeospira Circeis Eulaeospira Helicosiphon Janua subgenus Genus Paralaeospira Pileolaria subgenus
Duplicarla Pileolaria Protoleodora Simplicaria Dextralia Protolaeospira Bushiella Romanchella Spirorbella Spirorbis
Dexiospira Fauveldora Janua Leodora PUospira
Prodexiospira Protolaeospira subgenus Romanchella subgenus Spirorbis subgenus
Metalaeospira Neomicrorbis Paradexiospira
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anterior peg. Abdominal setae with vestigial brushtop. Incubation in faecal groove on attachment stalk. Duplicara Vine 1972, Spirorbis (Laeospira) koehleri Caullery and Mesnil 1897; 3 species. Sinistraliy coiled; two pairs of thoracic tori present. Collar setae of fm and blade construction, not crossstriated. Side setae present; thoracic uncini with several rows of teeth. Anterior abdominal tori divided into two parts. Operculum with two or more plates stacked on top of each other; incubation in the operculum. Eulaeospira Pillai 1970, Spirorbis (Laeospira) orientalis Pillai 1960; 2 species. Sinistraliy coiled; two pairs of thoracic tori. Collar setae of fin and blade construction, without crossstriations. Side setae absent. Uncini with teeth in several rows; anterior peg bifid or multifid. Incubation in groove along posterior abdomen. Fauveldora Knight-Jones 1972, Jatiua (Fauveldora) kayi Knight-Jones 1972; only species. Sinistraliy coiled; two pairs of thoracic tori present; collar setae limbate; abdominal setae with blades as big as those in the collar setae; side setae in third fascicle present. Uncini with teeth in several rows. Incubation in the operculum. Helicosiphon Gravier 1907, H. biscoeensis Gravier 1907; only species. Sinistraliy coiled, or almost uncoiled; three thoracic setigers on concave side. Collar setae limbate; side setae absent. Thoracic uncini with blunt anterior peg. Abdominal tori distinctly larger on concave side. Incubation in faecal groove attached by stalk. Janua Saint-Joseph 1894, Spirorbis pagenstecheri Quatrefages 1865; 7 species. Dextrally coiled; two pairs of thoracic tori present; collar setae limbate; side setae in third fascicle present. Abdominal setae with blades as big as those in the collar seate. Uncini with teeth in several rows. Incubation in the operculum. Leodora Saint-Joseph 1894, Spirorbis laevis Quatrefages 1865; 8 species. Sinistraliy coiled; two pairs of thoracic tori present; collar setae limbate, not dentate; side setae absent. Abdominal setae with blades as big as those in the collar setae. Uncini with teeth in several rows. Incubation in the operculum. Metalaeospira Pillai 1970, Spirorbis pixelli Harris 1969; 4 species. Sinistraliy coiled; three thoracic tori on the concave side. Collar setae with limbate blades; side setae present. Both thoracic and abdominal uncini more numerous on concave side. Thoracic uncini with blunt
Amplaria Knight-Jones 1973, A. spiculosa KnightJones 1973; only species. Sinistraliy coiling; four thoracic tori and four fascicles of setae on the concave side; three thoracic tori and five fascicles of setae on the convex side. Fin and blade collar setae without cross-striations; side setae in third and fourth thoracic fascicle. Thoracic uncini with blunt anterior peg; abdominal setae genie ulate, less than one fourth the size of the side setae. Abdomen somewhat asymmetrical; incubation in the operculum. Anomalorbis Vine 1972, A. manuatus Vine 1972; only species. Probably dextrally coiling; four complete thoracic setigers and remnants of a fifth. Collar setae limbate; side setae absent. Pronounced abdominal asymmetry; incubation probably in tube. Bushiella Knight-Jones 1973, Spirorbis evolutus Bush 1904b; 4 species. Sinistraliy coiling; two pairs of thoracic tori present; collar setae limbate, numerous. Side setae present in third fascicle. Thoracic uncini with blunt anterior peg. Margin of collar unfused; abdomen not strongly asymmetrical. Incubation in faecal groove with attachment stalk for the embryos. Capeospira Pillai 1970, Spirorbis (Paralaeospira) adeonella Day 1963; only species. Sinistraliy coiling; thorax with three complete and a fourth rudimentary setiger; collar setae of fin and blade construction; side setae absent. Incubation in the tube. Circeis Saint-Joseph 1894, Serpula spirillum Linnaeus 1758; 3 species. Dextrally coiled; three thoracic setigers present; collar setae limbate. Side setae absent. Incubation in the tube. Dexiospira Caullery and Mesnil 1897, Serpula corrgala Montagu 1803; 23 species. Usually dextrally coiled; two pairs of thoracic tori present. Collar setae limbate; abdominal setae with blades as long as those in the collar seta; side setae absent. Margins of collar fused to form dorsal tunnel; incubation in operculum. Dextralia Knight-Jones 1973, Spirorbis (Paradexiospira) falklandica Pixell 1913a; only species. Dextrally coiled; three rows of tori on the concave side and traces of a fourth on the convex side. Collar setae of fin and blade construction; side setae in third and fourth fascicle. Thoracic uncini with bilobed
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51
anterior peg. Incubation in faecal groove; attachment stalk absent. Neomicrorbis Roverto 1904, Serputa crenatostriatus Goldfuss 1833; 3 species. Sinistrally or dextraliy coiled; approximately seven thoracic setigers. Collar setae of fin and blade construction; side setae present in posterior thoracic setigers from the third. Paradexiospira Caullery and Mesnil 1897, Serpula vitrea Fabricius 1780; 3 species, Dextraliy colled; three complete and one fourth incomplete thoracic seliger. Collar setae of fin and blade construction; side setae present in the third fascicle. Incubation in the tube. Paralaeospira Caullery and Mesnil 1897, Spirorbis (Paralaeospira) aggregata Caullery and Mesnil 1897; 8 species. Sinistrally coiled; three thoracic tori on the concave side; collar setae of fin and blade construction without cross-striations; side setae present in the third fascicle. Thoracic and abdominal uncini more numerous on the concave side; thoracic uncini with blunt anterior pegs. Incubation in faecal groove; attachment stalk absent. Pileolaria Claparde 1870a, P. militaris Claparde 1870a; 20 species. Sinistrally coiled; two pairs of thoracic tori present; collar setae of fin and blade construction; blades usually coarsely dentate, cross-striated. Side setae present. Incubation in operculum. Pillaiospira Knight-Jones 1973, Janua (Pillaospira) trifurcata Knight-Jones 1973; only species. Dextraliy coiled; two pairs of thoracic tori; collar setae limbate, abdominal setae with blades as long as those in the collar setae; side setae absent. Incubation in the operculum. Prodexiospira Pillai 1970, Spirorbis violaceus Levinsen 1884; only species. Dextraliy coiled; three complete and a fourth incomplete seliger present. Collar setae limbate; side setae present in the third fascicle. Incubation in the tube. Protolaeospira Pixell 1912, P. ambilateralis Pixell 1912; 7 species. Sinistrally coiled; three pairs of tori and in addition a fourth incomplete present. Collar setae of fin and blade construction; side setae in third and fourth fascicle. Thoracic uncini with bilobed anterior pegs. Incubation in faecal groove; attachment stalk present. Protoleodora Pillai 1970, Spirorbis asperatus Bush 1904b; only species. Sinistrally coiled; three (two?) setigers in the thorax;
collar setae limbate; side setae present. Incubation in operculum. Romanchella Caullery and Mesnil 1897, R. perrieri Caullery and Mesnil 1897; 4 species. Sinistrally coiled; two pairs of thoracic tori present; collar setae limbate, few in number. Side setae in third fascicle. Thoracic uncini with blunt, gouge-shaped pegs. Margin of collar fused dorsally. Incubation in faecal groove, attachment stalk present. Simplicaria Knight-Jones, 1973, Spirorbis (Laeospira) pseudomiliiaris Thiriot-Quievreux, 1965; 3 species. Sinistrally coiled; two pairs of thoracic tori present; collar setae of fm and blade construction; blades usually coarsely dentate, cross-striated. Side setae absent. Incubation in operculum. Spirorbella Chamberiin 1919, Spirorbis (Dexiospira) marioni Caullery and Mesnil 1897; 3 species. Dextraliy coiled; two pairs of thoracic tori; collar setae of fin and blade construction; side setae in third fascicle. Thoracic uncini with broad anterior pegs. Incubation in tube, attachment stalk absent. Spirorbis Daudin 1800, Spirorbis borealis Daudin 1800; 48 species. Sinistrally coiled; thorax with two pairs of tori; collar setae of fin and blade construction; side setae present in the third fascicle. Thoracic uncini with broad anterior pegs. Incubation in the tube, attachment stalk absent. Taxonomic Notes Vine (1972) pointed out that the coiling of the spirorbids may not be a character of particularly high value and that even within one species, the coiling may vary from sinistral to dextral. It is thus of the utmost importance that great care be taken in using the key and definitions given above and that all the important features be checked carefully. It is quite possible that the coiling will turn out to be of great value in most of the genera, even if the character should turn out to be useless in certain instances. Invalid Genera Charybs Montfort in Mrch 1863, questionably Spirorbis Coretus Baster 1762, see Circeis Dexiorbis Chamberiin 1919c, see Dexiospira Heterodisca Fleming 1825, indeterminable Laeospira Caullery and Mesnil 1897, set Spirorbis Marsipospira Bailey 1969, see Protolaeospira Mera Saint-Joseph 1894, see Janua Neodexiospira Pillai 1970, see. Dexiospira
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Pixellia Pillai 1970, see Proiolaeospira Sinistrella Chamberlin 1919c, see Leodora Spirorbides Chamberlin 1919c, see Paradexiospira Stoa Serres 1855, indeterminable
FIVE "ARCHIANNELIDAN" FAMILIES
consider them free-standing families of the Polychaeta. Like Jouin and Westheide, I find it difficult to associate the five families, separately or as a unit, to any known polychaete family, and have for that reason listed them below, in alphabetical order. I do not think the five families are particularly closely related to each other. 1932
The following five families usually are considered as belonging to a separate order, Archiannelida. They are characterized, as pointed out by Hermans (1969), by features adaptive to life in the mesopsammon, and the features in which they resemble each other are exactly those that have been considered to be of selective advantage for the mesopsammon (Swedmark 1964; Hermans 1969). In agreement with Jouin (1971) and Westheide (1971), I find it difficult to retain the concept of the Archiannelida (Fauchald 1974a) and presently
FAMILY DINOPHILIDAE REMANE
Very small, usually distincdy segmented polychaetes, without appendages, setae and parapodia; some forms with adhesive glands posteriorly. Ciliary bands on ventral side present, and, usually segraentally arranged, ciliary tracts present. The family recently was reviewed by Jouin (1971) and by Westheide (1971).
Key to Genera la. 1 b. 2a (lb). 2b (lb). 3a (2b). 3b (2b). Muscular pharynx absent Muscular pharynx present Body slender and elongated; prostomium trilobed Body short, more or less inflated; prostomium rounded Pygidium with two adhesive glands; eyes absent Pygidium conical, without adhesive glands; eyes usually present Apharyngtus 2 Trilobodrilus 3 Diurodrilus Dinophilus
Generic Definitions Apharyngtus Westheide 1971, A. punicus Westheide 1971; only species. Slender, distinctly segmented bodies; pharyngeal apparatus absent; in addition to the regularly arranged ciliary bands, very fine cuticular hairs covering the body. Dinophilus Schmidt 1848, D. vorticoides Schmidt 1848; 8 species. Short-bodied, indistinctly segment dinophilids; pygidium rounded or conical. Pharyngeal apparatus present and well developed. Some forms distinctly sexually dimorphic. Diurodrilus Remane 1925, D. minimus Remane 1925; 5 species. Body short, inflated and indistinctly segmented. Pygidium with paired adhesive glands ending distally
on conical projections of various shapes. Pharyngeal apparatus well developed. Trilobodrilus Remane 1925, T. heideri Remane 1925; 3 species. Body long and slender; prostomium trilobed; eyes absent; pygidium rounded or conical, without adhesive glands. Pharyngeal apparatus well developed.
FAMILY NERILLIDAE LEVINSEN
1883
Interstitial small forms with few segments. Prostomium with paired palps, nuchal organs and up to three antennae. Ventral eversible pharynx with four jaws or unarmed. Uniramous parapodia with capillary setae or composite setae in a single fascicle. Parapodial cirri usually present. The key to genera and definitions given below have been modified after Jouin (1971). The main changes involve terminology to fit the terms used in this paper.
Key to Genera la. lb. 2a (la). 2b (la). 3a (2b). 3b (2b). Body with nine segments including the post-peristomial pharyngeal segment Body with less than nine segments Composite setae present Simple capillary setae present Three antennae present Maximally one antenna present 2 4 Mesonerilla 3 , Nerilla Meganerilla
7976
153
FIGURE 39. (A), Family DINOPHILIDAE, Diurodhlus ankeli, from Jouin, 1971, about 50x; (B), Family POLYGORDIIDAE, Polygordius neapotitanus, 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, Mesonerilta sp., after Jouin, 1971, about 25x.
i54 4a 4b 5a 5b 6a 6b 7a 7b 8a 8b (lb). (lb). (4a). (4a). (5a). (5a). (6b). (6b). (4b). (4b).
NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY Eight segments, including the post-peristomial pharyngeal segment Seven segments present , Both simple and composite setae present All setae either composite or simple All setae composite All setae simple and capillary Pharyngeal segment asetigerous, antennae absent; freshwater Pharyngeal segment with setae, two antennae present; marine Composite setae; parapodial cirri absent, palps reduced Setae simple and capillary; parapodial cirri present except on last segment
Science Series 28 5 g Nerillidopsis 6 Thaiassochaetus 7 Troglochaetus Nerillidium Paranerilla Psammoriedlia
Generic Definitions Meganerilla Boaden 1961, M. swedmarki Boaden 1961; 2 species. Body with nine segments including the pharyngeal segment. Two large palps and maximally one antenna present. All setae simple capillaries. Mesonerilla Remane 1949, M. luederitzi Remane 1949; 6 species. Body with nine segments including the pharyngeal segment. Two palps and three antennae present. All setae composite. Nerilla Schmidt 1848, N. antennata Schmidt 1848; 7 species. Body with nine segments including the pharyngeal segment. Three antennae and two palps present. All setae simple capillaries. Nerillidium Remane 1925, N. gracile Remane 1925; 5 species. Body with eight segments including the pharyngeal segment. Pharyngeal segment with setae; maximally two antennae present, may be absent. All setae simple capillaries. Nerillidopsis Jouin 1967, A', hyalina Jouin 1967; only species. Body with eight segments including the pharyngeal segment. Anterior and posterior setae simple capillaries; intermediate segments with composite setae. Paranerilla Jouin and Swedmark 1965, P. limicola Jouin and Swedmark 1965; only species.
Body with seven segments including the pharyngeal segment. Antennae and parapodial cirri absent; palps reduced to triangular lateral comers on the prostomium. Mud-dwelling forms. Setae composite. Psammoriedlia Kirsteuer 1966, P. ruperti Kirsteuer 1966; only species. Body with seven segments including the pharyngeal segment. Antennae absent; two palps present; parapodial cirri present, except on last segments. Setae simple capillaries. Thaiassochaetus Ax 1954, T. palpifoliaceus Ax 1954; only species. Body with eight segments including the pharyngeal segment. Antennae and parapodial cirri absent; palps present. All setae composite spinigers. Troglochaetus Delachaux 1921, T. beranecki Delachaux 1921; only species. Body with eight segments, including the pharyngeal segment. Antennae absent; palps present; pharyngeal segment asetigerous. All setae simple capillaries. Freshwater and caves.
FAMILY POLYGORDIIDAE CZERNIAVSKY
1881a
Slender, mainly interstitial forms. Prostomium with two solid antennae and nuchal slits. Eversible or noneversible muscular pharynx ventral. Eyes and parapodia absent; setae absent in most forms; segmentation and ciliation poorly developed.
Key to Genera la. lb. Setae present in posterior segments Setae absent Generic Definitions Chaetogordius Moore 1904a, C. canaliculatus Moore 1904a; only species. Chaetogordius Polygordius Prostomium conical with paired, slightly articulated antennae and large nuchal organs. Segmentation indistinct anteriorly, more distinct posteriorly. Single slender capillary setae in each of the last 10-12 seg-
7976
THE POLYCHAETE WORMS FAMILY PROTODRILIDAE CZERNIAVSKY
155 1881a
ments; only one segment with paired setae on either side; all others with a single seta on either side. Polygordius Schneider 1868, P. lacteus Schneider 1868; 15 species. Prostoitiium rounded or conical with paired, usually smooth antennae and large nuchal organs. Segmentation usually indistinct along the whole body. Posterior end of body usually slightly wider than the rest of the body. Setae absent.
Interstitial slender forms. Prostomium with paired, hollow or solid antennae; eyes present. Pharynx muscular, but not eversible. Segmentation and parapodia poorly developed, setae absent in most known species; parapodia always absent. The family has been reviewed recently by Jouin (1966).
Key to Genera la. lb. Setae or segmentally arranged adhesive organs present Setae and segmentally arranged adhesive organs absent Protodriloides Protodrilus 1881a
Generic Definitions Protodriloides Jouin 1966, Protodrilus chaetifer Remane 1926; 2 species. Salivary glands and lateral organs absent; antennae solid. Setae or segmentally arranged adhesive glands present. Protodrilus Czerniavsky 1881a, P. mirabilis Czerniavsky 1881a; 20 species. Salivary glands and lateral organs present; antennae hollow. Setae and segmentally arranged adhesive glands absent.
FAMILY SACCOCIRRIDAE CZERNIAVSKY
Interstitial slender forms. Prostomium with paired hollow antennae; eyes present. Proboscis a ventral muscular pad, present in most forms, but may be absent. Parapodia uniramous, with simple, chiselshaped setae. The family is known for a single genus, Saccocirrus Bobretzky 1872, with genotype, S. papillocercus Bobretzky 1872 and about 12 species. Jouin (1971) gave a key to species.
156
NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY GLOSSARY
Science Series 28
The glossary does not contain terms in general use in zoology or in biology in general, such as distal and proximal, anterior and posterior, etc. Generally, these terms can be found in a good dictionary. The glossary contains the terms characteristically used in systematic polychaete literature and explanation of idiosyncratic usage where terms are used in a different sense from the usual lexicographic one. Number in parenthesis refer to the specific example of this feature illustrated in the appended illustrations.
branchia (e)any extension of the body wall with a loop of the vascular system or which is wellequipped with capillary blood-vessels; a gill (27). buccalpertaining to the mouth.
capillary (ies)in polychaete literature used either as an adjective in the combination capillary seta or as a noun characterizing long, slender tapering setae. caruncleposterior sensory organ projecting from the prostomium sometimes over the first several segments (28). cephalonhead. cephalic cagelong protective setae enclosing and protecting the head. cephalic rimflange encircling the head (25). cephalic veilhoodlike membrane between the paleae and the buccal tentacles in pectinariids. ceratophorebasal joini of an antenna (30). ceratostyledistal joint of an antenna (29). chevronV-shaped chitinized jaw piece at the base of the eversible pharynx in some goniadids (31). cirriformslender and cylindrical (41). cirrophorebasal joint of a cirrus (35). cirrostyledistal joint of a cirrus (34), cirrus (1)sensory projection, usually slender and cylindrical, from the superior part of the notopodiura (dorsal cirrus) or from the inferior part of the neuropodium (ventral cirrus) (36,38,78,80). clvateclub-shaped (32). companion setaesmall, simple setae in rows, accompanying, or alternating with larger setae, usually hooks of some kind. compound (or composite) setae^jointed setae (37).
abdomenposterior part of the body, behind the thorax, sometimes followed by a tail (2). aclculiun (a)stout supportive setae found internally in each parapodial ramus where these project from the body. Acicular setae are thick, projecting setae (5). aileronaccessory jaw plate in the glycerids (9), anal cirrus (i)one or more elongated projections from the pygidium (11). antenna (e)sensory projection arising from the dorsal, lateral or frontal surface of the prostomium; innervated from the first part of the brain (12,61). apodouswithout parapodia (66). aristateabout setae: simple setae with smooth shaft and a tuft of hairs or a single spine distally (17), arborescentbranching (like a tree) (18). asetigerouswithout setae (66). auricularear-shaped (19). avicularbeaked (shaped like a bird's head) (20).
biarticulatewith two joints; used about antennae, tentacles and palps (13). bidentatewith two teeth (21). bifidsplit in two (22). bifurcatewith two prongs (23). bilimbatesimple setae with two wings or flattened margins (24). bipinnateformed like a feather with a main stem and two rows of side branches (26). biramouswith two branches; usually used about parapodia having both noto- and neuropodia present (6).
deciduousliable to fall off (like leaves). dentatewith teeth (39). denticulatewith small teeth (40). digitiformfinger-shaped (42).
elytron (a)dorsal scales found in the aphroditoids; homologous with the dorsal cirrostyles (43).
1976
157
elytrophoreaphroditoid cirrophore carrying an elytron (44). epitokemodified reproductive stage or specimen, often swarming.
facial tubercleprojecting ridge or lobe on the upper lip of certain polychaetes (especially scaleworms). falcatedistally curved, usually blunt (used about setae) (45). falcigerdistally blunt and curved setae (45). filifonnslender and threadlike (46). foUaceousleaflike (47). fusiformcigar- or spindle-shaped (48).
geniculatedbent (like a knee) (49),
harpoon setastout pointed seta with recurved barbs near the tip (50). liemigomph articulationwith asymmetrical articulation nearly at right angles to the long axis of the shaft (52). heterogomph articulationwith articulation clearly oblique to the long axis of the shaft (53). homogomph articulationwith articulation distinctly and symmetrically at right angles to the long axis of the shaft (54). hooded setaesetae distally covered by a delicate chitinous envelope or guard (51). hookgeneral term used about stout-shafted, blunt, often distally curved and dentate setae; smaller hooks arranged in single or double rows are often referred to as uncini. imbricatedoverlapping (like tiles). inferiorthe more ventral (of two or more structures). ligulefinger-shaped major process on a parapodium (55). limbatesimple seta with a flattened margin or wing (56). lobemajor parapodial process, used mainly about flattened kinds, but also more generally about all kinds of major parapodial processes (8).
FIGURE
40. Diagram of selected terms defined in the Glossary
#1-18.
158
Science Series 28
52
-53
-54
FIGURE
41. Diagram of selected terms defined in the Glossary #19-60.
]976
THE POLVCHAETE WORMS
159
long-handledused about uncini, with a long basal rod supporting the uncinus
metastomiumrarely used terin covering the segmented body between the prostomium and pygidium, but including neither. moniliformbeaded or beadlike (57). muerowith a sharp point or tip; abruptly tapered (58). muitiarticulatedwith several joints (60).
natatoryswimming. neuropodiuin (a)ventral branch (ramus) of a parapodium (7). neurosetaesetae of the neuropodium. notopodium (a)dorsal branch (ramus) of a parapodium (3). ^,^.|,^ ^.^^^ notosetaesetae of a notopodium. nuchalpertaining 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.
occipitalpertaining to the posterodorsal part of the prostomium (64). ocularpertaining to the eyes. ocular peduncleprojecting structure supporting the eyes, especially in the polyodontids. onunatophoresee ocular peduncle. operculum (a)literally a lid; used about a structure some tubicolous worms use as stoppers for their tubes when the occupant is retracted; see Sabellariidae and Serpulidae.
palea (e)strong or broad, usually flattened setae. palmateresembling the fronds of a palm (67). palp (s)sensory or feeding structures innervated from the posterior part of the brain or from the circumesophagal nerve ring; anteroventral and sensory in the Phyllodocida; posterodorsal and used in feeding in the spioniform worms (13). 42. Diagram of selected terms defined in the Glossary #61-83.
FIGURE
paragnathchitinous denticle in the pharyngeal cavity of nereids (63).
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Science Series 28
parapodium (a)segmentaily arranged projections carrying setae; foot (16). pectinatecombiike; with series of projections arranged like the teeth of a comb (68), penicillatebrushlike (70). peristomium (a)first distinct post-prostomial region; strictly including only the region around the tnouth, in practice including also segments fused to this structure, forming the posterior part of the recognizable head (16,66). pharynx (ges)anterior part of the digestive tract; often eversible, always modified for feeding purposes, sometimes also for burrowing (62). pinnatefeatherlike, with a main stem and lateral branches, see also bipinnate (71). plumoseresembling down; hairy (69). postsetalposterior to the setae; used about parapodial lobes or hgules (8). presetalanterior to the setae; used about parapodial lobes or ligules (4). prostomium (a)anteriormost, a pre-segmental part of the body anterior to the mouth, enclosing at least the anterior part of the brain, often with antennae and eyes (14,64). proventriclemuscularized region of the anterior digestive tract in syllids, found posterior to the pharynx. pygidium (a)post-segmental terminal part of the body carrying the anus (10).
setasecretion from the parapodia forming the armature of these structures. settersegment carrying setae. short-handledused about uncini, without a long rod-shaped support, fine threads may be present, simple setaunjointed seta. spatulateblade-shaped, usually blunt-tipped, sometimes with a muero (59). spinigerseta that tapers to a fine point; most frequently used about composite setae (33). spinous pocketenlarged, pocketlike serration along the margin of the setae of some scale-worms (74). stylodesmall, fingerlike projection associated with a parapodium, usually small and distinctly longer than wide (73). sub-biramouspertaining to parapodia in which the neuropodia are well developed and the notopodia reduced (77), subulateawl-shaped; tapering to a fine point (75), superiorthe more dorsal (of two or more structures).
tentacular cirrussensory projections arising either from the peristomium or from cephalized segments, in the latter case considered homologous with the dorsal and ventral cirri of normai, p--sicephalic parapodia (15,65). thoraxanterior region of the body, posterior to the head(l). trepanchitinized, anteriorly toothed part of the eversible pharynx of some worms, especially syllids (81). tridentatewith three teeth (82).
radicleone of the main tentacles in the tentacular crown of sabellids and serpulids. ramosebranched. reniformkidney-shaped (76), rugoseroughened, lumpy (72).
scapheflattened caudal appendage of pectinariids. secondary tooththe second of two teeth, the first being the apical, terminal or primary one. segmentany part of the body, apart from the prostomium and pygidium set off internally or externally by septa or otherwise from the preceding and following parts.
uncinus (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 transverse to the long axis of the animal (83). uneinigersegment carrying uncini. unidentatedistally entire (45). uniramouswith one branch only; used about parapodia in which one ramus, most frequently the notopodium, is absent (79).
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]976 LITERATURE CITED

ABSALON, K. AND S. HRABE.
\6l
1930. ber einen neuen Ssswasser-Polychaelen aus den Hhlengewssem der Herzegowina. Zool. Anz. 88:249-264. AiVAR, R. G. AND K. H. ALIKUNHI. 1940. On a new pisionid from Ihe sandy beach, Madras. Indian Mus, Calcutta, Rec. 42:89-107. 1943. Change of Ihe generic name Pisionella Aiyar and Alikunhi, 1940 into Pisionidens (Polychaeta). Current Science, Bangalore 12(4): 120. tvESSON, B. 1963. The comparative morphology and embryology of the head in scale worms (Aphroditidae, Polychaeta). Ark. Zool. Stockholm 16(7):125-163. 1967. The embryology of the polychaete Eunice kobiensis. Acta Zool. 48:141-192. ALIKUNHI, K.H. 1949. On Anophthalmus (Fam.: Hesionidae) a new genus of polychaetes with description of four new species from the sandy beach at Madras. Proc. 35th Indian Sei. Congr. p. 192.
ANDREWS, E.A.
1932. Calamyzas amphictenicola, ein ektoparasitischer Verwandter der Sylliden. Zool. Bidr. Uppsala 14:153-218. ASHWORTH, J.H. 1901. The anatomy oScalibregma inflatum Rathke. Quar. J. Micro. Sei., London 45:237-309. AuDOuiN, J.V. AND H. MILNE EDWARDS. 1830. Description de VHipponoe, nouveau genred'Annelides. Ann. Sei. Nat. Paris (1)20:156-159. -. 1832. Classication des Annelides, et description de celles qui habitent les ctes de la France. Ann. Sei. Nat. Paris (1) 27:337-347. 1833a. Classification des Annlides, et description de celles qui habitent les ctes de la France. Ann. Sei. Nat. Paris (1)28:187-247. 1833b, Classification des Annlides, et description de celles qui habitent les ctes de la France. Ann. Sei. Nat. Paris (1)29:195-269. 1833c. Classification des Annlides, et description de celles qui habitent les ctes de la France. Ann. Sei. Nat. Paris (1) 29:388-412. 1833d. Classication des Annlides, et description de celles qui habitent les ctes de la France. Ann. Sei. Nat. Paris (1) 30:411-425. 1834. Recherches pour servir l'histoire natitfelle du littoral de la France, ou Receuil de mmoires sur l'anatomie, la physiologie, la classification et les moeurs des animaux de nos ctes; ouvrage accompagn des planches faites de aprs nature. 2. Annlides, l'pt. Paris, 290 pp. AucENER, H. 1913a. Polychaeta I, Errantia. In: Michaelsen, W. and R. Hartmeyer (eds.): Die Fauna SdwestAustraliens. Ergebnisse der Hamburger sdwest-australischen Forschungsreise 1905. 4(5):65-304. 1913b. Polychaeten von Franz Josephland II. Zool. Anz. 41:253-273. 1914. Polychaeta 11, Sedentaria. In: Michaelsen, W. and R. Hartmeyer (eds.): Die Fauna SdwestAustraliens. Ergebnisse der Hamburger sdwest-australiscben Forschungsreise 1905. 5(I):I-170. 1918. Polychaeta. n: Michaelsen, W. (ed.): Beitrge zur Kenntnis des Meeresfauna West-Afrikas. 2(2):67-625. 1922. ber littoral Polychaeten von Westindien. Sitzber. Ges. Naturf. Freunde Beriin, 1922:38-63. 1926. Polychaeten von Neuseeland II. Sedentaria. Vidensk. Medd. dansk naturh. Foren. 81:157-294. 1927a. Polychaeten von Sdost- und SdAustralien. Vidensk. Medd. dansk naturh. Foren. 83: 71-275. 1927b. Polychaeten von Curacao. Bijdragen tot de Kennis der Fauna van Curacao. Resultaten eener Reis van Dr. C.J. van der Horst in 1920. Zool. Genoots. "Natura aitis Magistra", Amsterdam 25:39-82. 1928a. Beitrag zur Polychaetenfauna der Ostsee. Zeits. Moiph. Oekol. Tiere 11:102-104. .. 1928b. Ergnzung zu den Polychaeten von Spitzbergen. Fauna rctica, Jena 5:647-834. Ax, P. 1954. Thalassochaetus palpifoliaceus, nov. gen., nov. spec, (Archiannelida, Nerillidae), ein marine Verwandtes von Troglochaetus beranecki Delachaux. Zool. Anz. 153: 64-75.
1891. Report upon the Annehda Polychaeta of Beaufort, North Carolina. U.S. Nat. Mus., Proc. 14:277-302. 1925. Beitrge zur Kenntnis die PolychaeteFauna Russlands. C.R. Acad. Sei. SSSR, Leningrad, 1925:125-126. 1927. ber die pontokaspischen Polychaeten. I, Die Gattungen Hypania Ostrooumov und Hypaniola n. gen. Ann. Mus. Zool. Leningrad 28:48-62.
ANNENKOVA, N.
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1881. New England Annelida. Pt. I. Historical sketch, with annotated list of the species hitherto recorded. Trans. Conn. Acad. Arts Sei. 4:285-324. 1885. Notice of recent additions to the marine Invertebrata of the northeastern coast of America, with description of new genera and species and critical remarks on others. U.S. Nat. Mus., Proc. 8:424-448, 1900. Additions to the Turbellaria, Nemertina and Annelida of the Bermudas, with revisions of some New England genera and species. Trans. Conn. Acad. Arts Sei. 10:595-671. VicuiER, C. 1886. tudes sur les animaux infrieurs de la Baie d'Alger. Recherches sur les Annlides plagiques. Arch. zool. exp. gen. Paris (2) 4:347-442. 1911. Nouvelles tudes sur le plancton de la Baie d'Alger. Ann, Sei. Nat. Paris (9) 13:187-267. 1912. Nouvelles tudes sur le plancton de la Baie d'Alger. Trois Aphroditiens plagiques. Ann, Sei, Nat. Paris (9) 15:85-99.
VINE, P.J.
[943. Biological results of the last cruise of the CARNEGIE. Polychaetous annelids. Sei. Res. Cruise VII CARNEGIE 1928-1929:30-59. UcHlDA, H. 1968. Polychaetous annelids from Shakotan (Hokkaido) The collection in 1967. J. Hokkaido Univ, Fac. Sei. (6) Zool. 19:595-612. UUANiN, B. 1878. Sur le genre Sagitella N. Wagner. Arch, zool. exp. gen. Paris 7:1-32. UscHAKOV, P. 1953. [New species of polychaetous annelids of the family Phyllodocidae (Polychaeta)]. (In Russian). Trudy Zool. Inst. Akad. Nauk SSSR Leningrad 13: 207-209. 1955a. [Polychaetous annelids of the Far Eastern Seas of the USSR]. (In Russian). Akad. Nauk SSSR, Keys to the Fauna of the SSSR 56:1-433 (translated, 1965 by Israel Program Scientific Translating, Jerusalem). 1955b. [Polychaetous annelids of the family Aphroditidae from the Kurilo-Kamshatska Trench]. (In Russian). Trudy Inst. Ocean. Akad. Nauk SSSR 12: 311-321. 1972. [Polychaeta 1. Polychaetes of the suborder Phyllodociformia of the Polar Basin and the northwestern part of the Pacific]. (In Russian). Akad. Nauk SSSR. Zool. Inst. Fauna of the SSSR, n.s. 102, 271 pp. (translated 1974 by the Israel Program for Scientific Translation, Jerusalem), AND B.L. Wu. 1959. [The polychaetous annelids of the families Phyllodocidae and Aphroditidae from the Yellow Sea]. (In Chinese and Russian). Arch. Inst. Oceanol. Sinica 1(4): 1^0. 1890. Histoire naturelle des Annels marins et d'eau douce. Lombriciens, Hirudiniens, Bdellomorphs, Trtulariens et Planariens. 3(2):340-768. Paris, Roret. VEJDOVSKY, F. 1884. System und Morphologie des Oligochaeten. Prague. 166 pp. VERRILL, A.E. 1871. Marine fauna of Eastport, Maine. Bull. Essex Inst. 3:2-6. 1873a. Results of recent dredging expeditions on the coast of New England. Amer. J. Sei. New Haven (3) 5:1-16. 1873b. Report upon the invertebrate animals of Vineyard Sound and the adjacent waters, with an account of the physical characters of the region. Rep. U.S. Comm. Fish. 1871-72:295-778. 1874. Explorations of Casco Bay by the U.S. Fisheries Commission, in 1873. Proc. Amer. Ass. Adv. Sei. Salem, Mass. 22:340-395. 1880. Notices of recent additions to the marine Invertebrata of the northeastern coast of America, with descriptions of new genera and species and critical remarks on others. Pt. 1. Annelida, Gephyrea, Nemertina, Nematoda, Polyzoa, Tunicata, Mollusca, Anthozoa, Echinodermata, Porifera. U.S. Nat. Mus., Proc. 2: 165-205,
VAILLANT, L.
1972. Spirorbinae (Polychaeta: Serpulidae) from the Red Sea including descriptions of a new genus and four new species. J. Linn. Soc. London Zool, 51:1-16.
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WEBSTER, H.E.
1879a. The Annelida Chaetopoda of New Jersey. Ann. Rep. N.Y. State Mus. Nat. Hist. 32: 101-128. _. 1879b. Annelida Chaetopoda of the Virginian coast. Trans. Albany Inst. N.Y. 9:202-269. 1880. Annelida Chaetopoda of New Jersey. Ann. Rep. N.Y. State Mus. Nat. Hist. 39:128-159 (reprint of the 1879 paper with added plates). . AND J.E. BENEDICT. 1884. The Annelida Chaetopoda from EVovincetown and WeUfleet Mass. Rep. U.S. Fish. Comm. 1881:699-747. -. 1887. The Annelida Chaetopoda from Eastport, Maine. Rep. U.S. Fish. Comm. 1885:707-755.
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1959. The genera of Arenicolidae (Polychaeta). Proc, Zool. Soc. London 133:301-314.
WESENBERG-LUND, E.
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1971. Apharyngtus punicus nov. gen. nov. spec., ein aberranter Archiannelide aus dem Mesopsammal der tunesischen Mittelmeerkste. Mikrofauna des
1976
179
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1851. Report on the British Annelida. Rep. Brit. Assoc. Adv. Sei. London 21:159-272.
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1935. ber das Vorkommen der Brackwasser-poiychaete Manayunkia (M. polaris n. sp.) an der Murmankiiste. Zool. Anz. 109:195-203. ZEPPELIN, M. VON. 1883. ber den Bau und die Teilungsvorgnge des Ctenodrilus monostylos n. sp. Zeits. wiss. Zool, 39:615-652.
ZiBROwius, H. 1969. Review of some little known genera of Serpulidae (Annelida: Polychaeta). Smithsonian Contr. Zool. 112:1-22. Accepted for publication June 30, 1975.
180
NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY INDEX Page numbers set in bold face type represent citations of illustrations. Amphitritides, 129, 130 Amphitritoides, 133 Amphytrite, 133 Ampiara 149, 150 Amythas, 123, 125 Amythasides, 123, 125 Amytis, 85 Anaitides, 48 Anamobaea, 140 Anaspio, 23 Ancistargis, 78 Ancistria, 35 Ancistrosyllis, 78 Andresia, 56, 60 Andromache, 111 AnguillosylUs, 80, 81 Anisoceras, 113 Anisocirrus, 133 Anobothrella, 123, 125 Anobothrus, 124, 125 Anomaiorbis, 148, 149, 150 Anophthalmus, 77 Anoplonereis, 77 Anoplopisione, 73 Anoplosyllis, 85 Anotomastus, 33, 36 Anthostoma, 17 Antinoana, 56, 60 Antinoe, 57, 60 Antinoella, 57, 60 Antiobactrum, 42 Antonbruunia, 91 Antonbruunidae, 8, 12, 91 Aonides, 22, 23 Aonis, 25, 97 Aonopsis, 25 Aotearia, 109 Aparaonis, 17, 18 Apharyngtus, 152 Aphelothrix, 107 Aphlebina, 133 Aphrodita, 53, 54 Aphroditidae, 8, 10, 53-55, 54 Aphrogenia, 54 Apistobranchidae, 8, 14, 21, 22 Apistobranchus, 22 Apneuma, 133 Apomatopsis, 147 Apomatus, 141, 143 Aponobranchus, 135 Apophryotrocha, 112 Apoprionospio, 23 Aporosyllis, 85 Aquilaspio, 23 Arabella, 110, 111 ArabelUdae, 8, 13, 108, 110-111 Arabes, 111 Aracoda, 111 Archidice, 30 Arcteobia, 57, 60 Aretonoe, 59, 60 Arctonoella, 59, 60 Arenia, 35 Aicnicola, 37
Science Series 28
Abatenicola, 37 Aberranta, 25 Abyssoclymene, 39 Acanthicolepis, 58, 59 Aceionereis, 91 Acesta, 18 Acholoe, 59 Acicularia, 100 Acoetes, 66 AcfociiTidae, 8, 13. 14, 29, 30-31 Acrocirrus, 30 Admetella, 59 Adyte, 57, 60 Aedicira, 17, 18 Aenone, 111 Aglaophamus, 97 Aglaopheme, 97 Aglaura, 111 Aglaundes, 111 Agnodice, 65 Alcandra, 17 Alciopa, 51,52 Alciopidae, 8, 12, 46, 51-52 Alciopina, 51, 52 Alentia, 59, 60 Alentiana, 59, 60 Alcunhia, 73, 75 Alitta, 91 Alkmaria, 123, 124 Allia, 18 Allmaniella, 58, 60 Alluaudella, 79, 81 Aloysina, 43 Amaea, 133 Amaeana, 129, 130 Amage, 124 Amblyosyllis, 81 Ambo, 30 Amelinna, 121, 124 Americonuphis, 104, 105 Ammochares, 115 Ammotiypane, 43 Ammotrypanella, 42 Ampharana, 123, 124 Ampharete, 123, 124 Ampharetidae, 8, 10, 11, 121128, 122 Ampharetides, 134 Ampheiothrix, 107 Amphibranchus, 103 Amphicora, 140 Amphicorina, 140 Amphicteis, 124, 125 Amphictene, 120 Amphiduros, 75 Amphiglena, 137, 138 Amphinome, 100, 102 Amphinomidae, 8, 11, 100-103, 101 Amphiro, 107, 133 Amphiron, 107 Amphisamytha, 124, 125 Amphitrite, 129, 130
Arenicolidae, 8, 13, 31, 37 Arenicolides, 37 Areniella, 35 Arete, 91 Ariapithes, 120 Aricia, 17, 22 Aricidea, 17, 18 Aricideopsis, 25 Arippasa, 140 Aristenia, 117 Artnandia, 42 Artacama, 128, 130 Artacamella, 128, 130 Arwidssonia, 41 Aryandes, 128 Asabellides, 124, 125 Asclerocheilus, 44 Asloegia, 103 Aspe ira, 140 Asterope, 52 Asychis, 39 Atelesyllis, 80, 81 Athelepus, 133 Auchenoplax, 121, 125 Audouinia, 30 Augeneria, 109 Augeneriella, 137, 138 Australaugeneria, 56, 60 Auslralonereis, 87, 88 Austrolaenilla, 57, 60 Austrophyllum, 47, 48 Autolytides, 85 Autolytus, 79, 81 AutosylUs, 79, 81 Axionice, 130 Axiothea, 41 Axiothella, 39 B Baffmia. 129, 131 Balanochaeta, 117 BarantoUa, 32, 33, 36 Barbularia, 55 Barrukia, 57, 60 Bathya, 131, 133 Bathyadmetella, 59, 60 Bathychloeia, 100, 102 Bathyglycinde, 92, 93, 94 Bathymoorea, 58, 60 Bathynoe, 65 Bebryce, 98 Benhamipolynoe, 59, 61 Benthoscolex, 100, 102 Bergstroemia, 48, 49 Beikeleya, 15, 16 Bhawania, 72 Biborin, 110 Bilobaria, 25 Bispira, 137, 138 Blainvillea, 107 Blenda, 103 Boccaidia, 22, 24, 25 Boguea, 135
7976
Bogueidae, 8, 14, 122, 135 Boguella, 135 Bonhourelia, 142, 143 Bonuania, 74, 75 Bookhoutia, 93 Bouchiria, 58, 61 Brachysyllis, 85 Brada, 116 Bradabyssa, 116 Branchamphinome, 100, 102 Branchetus, 17 Branchioasychis, 39 Branchiocapitella, 32, 33, 36 Branchiomaldane, 37 Branchiomma, 137, 138, 140 Branchionereis, 91 Branchiosabella, 128 Branchiosyilis, 81 Branchoscolex, 35 Brandiika, 140 Brania, 80, 82 Branielia, 80, 82 Briaraea, 99 Bruunilla, 118 Bucheita, 35 Bushiella, 149, 150 Buskiella, 116 Bylgia, 65 Bylgides, 56, 61
18]
Cryptonota, 104 Cryptopomatus, 120 CtenodrUidae, 8, 13, 19, 19-20 Ctenodrilus, 19, 20 Ctenospio, 26 Cyanippa, 55 Cyaxares, 133 Cystonereis, 85 Cystopomatus, 147 D Dalhousia, 77 Dalhousiella, 75, 76 Danymene, 111 Dasybranchetus, 33, 33, 36 Dasybranchus, 33, 33, 36 Dasychone, 140 Dasychonopsis, 140 Dasylepis, 65 Dasymallus, 36 Dasynema, 142, 144 Dawbinia, 87, 88 Decamastus, 32, 34, 36 Dejoces, 133 Demonax, 137, 138 Dendrobranchus, 133 Dendronereides, 87, 88 Dendronereis, 86, 88 Dendrophora, 133 Desdemona, 137, 138 Desmosyllis, 85 Dexiorbis, 151 Dexiospira, 148, 149, 150 Dextralia, 140, 150 Dialychone, 137, 138 Didymobranchus, 103 Dilepidonotus, 58, 61 Dindymene, 43 Dindymenides, 42 Dinophilidae, 8, 9, 152, 153 Dinophilus, 152 Diopatra, 104, 105 Dioplosyllis, 80, 82 Diplobranchus, 97 Diploceraea, 85 Diplocirrus, 116, 117 Diplotis, 26 Dipolydora, 26 Dipomatus, 141, 144 Disoma, 26, 27 Disomides, 27 Dispio, 23, 24 Distylia, 140 Distylidia, 137, 138 Ditrocha, 36 Ditrupa, 142, 144 Diurodrilus, 152 Dodecaceria, 29, 30 Dorvillea, 112 Dorvilleidae, 8, 11, 12, 13, 111113, 112 Doyeria, 85 Drieschella, 59, 61 Drieschia, 58, 61 Drilognathus, 110 Drilonereis, 110, 111 Duplicaria, 149, 150
Cabira, 78 Caesicimis, 41 Calamyzas, 85 Calamyzidae, 8, 14, 74, 85 Calcareopomatus, 143 Califia, 16 Callizona, 52 Callizonella, 52 Campesyliis, 80, 82 Caobangia, 141 Caobangiidae, 8, 11, 136, 141 Capeospira, 149, 150 Capitella, 31, 32, 33, 36 Capitellethus, 32, 33, 36 Capiteilidae, 8, 13,31,31-36 Capitellides, 32, 33, 36 Capitita, 35 Capitobranchus, 33, 36 Capltomastus, 31, 32, 33, 36 Carazzta, 25 Carobia, 51 Cassandane, 43 Castalia, 77 CauUeriella, 29 Cenogenus, 109 Cenothrix, 111 Centrocorone, 120 Ceratocephale, 87, 88 Ceraionereis, 87, 88 Cervilia, 56, 61 Chaetacanthus, 58, 61 Chaetobranchus, 133 Chaetogordius, 154 Chaetoparia, 48, 49 Chaetopteridae, 8, II, 28, 28-29 Chaetoptenis, 28
Chaetosphaera, 25, 65 Chaetosyllis, 85 Chaetozone, 29 Chaponella, 41, 140 Chaiybs, 151 Chaunorhynchus, 91 CheUonereis, 86, 88 Cheironotus, 26 Cherusca, 27 Chinonereis, 91 Chitinopoma, 143 Chitinopomoides, 143, 144 Chloeia, 100, 102 Chloenea, 102, 103 Chloenopsis, 100, 102 Chloochaeta, 103 Chloraema, 117 Chone, 137, 138 Chorizobranchus, 37 Chrysodon, 120 Chiysopetalidae, 8, 10, 71, 71-72 Chrysopetalum, 72 Chrysothemis, 41 Circeis, 149, 150 Ciiraria, 51 Cirratulidae, 8, 13, 28, 29-30 Cirratulispio, 30 Cirratulus, 29 Cirrhineris, 30 Cimformia, 29, 30 Ciirobranchia, 111 Ciiroceros, 91 Cirrodoce, 47, 49 Cirronereis, 88, 91 Cirrophorus, 17, 18 Cirrosyllis, 77, 85 Cistena, 120 Cistenides, 120 Clapaiedia, 85 Clavadoce, 48, 49 Clavisyllis, 80, 82 Clavodorum, 98 Cleta, 52 Clymaldane, 39, 40 Clymene, 41 Ciymenella, 39, 40 Clymenides, 37 Ciymenopsis, 38, 40 Clymenura, 39, 40 Clytie, 17 Coelobranchus, 110 Colobranchus, 26 Colonianella, 103 Colymmatops, 129, 131 Commensodonim, 98 Conconia, 71 Conopomatus, 142, 144 Coppingeria, 116, 117 Corephorus, 135 Coretus, 151 Corynocephalus, 52 Cossura, 21 Cossuridae, 8, 13, 20, 21 Crithida, 85 Crosslandiella, 142, 144 Crossostoma, 128 Cnicigera, 142, 144 Cryptonereis, 87, 88
182 Dybowscella, 140 Dysponetus, 72 E
NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY Eimotomastus, 33, 34, 36 Eunotophyllum, 51 Eupallasia, 120 Eupanthalis, 66 Euphione, 58, 61 Euphionella, 58, 61 Eupholoe, 69, 70 Euphrosine, 103 Euphiosinidae, 8, II, 101, 103 Eupista, 133 Eupistella, 129, 131 Eupolymnia, 130, 131 Eupolynoe, 65 Eupolyodontes, 66 Eupomatus, 147 Eupompe, 66 Euratella, 137, 138 Eurato, 140 Eaiymedusa, 85 Eurysyllis, 80, 82 Eurythoe, 102 Eusamytha, 123, 125, 128 Emamythella, 123, 125 Euscione, 133 Euscleiocheilus, 45 Eusigalion, 71 Euspio, 26 Eusthenelais, 71 EusylUs, 80, 82 Euthalenessa, 69, 70 Euthelepus, 129, 131 Euzonus, 42, 43 Evarne, 65 Evamella, 65 Exallopus, 112 Exogone, 80, 82 Exogonella, 79, 82 Exogonita, 80, 82 Exogonoides, 80, 82 Exotokas, 85 Frennia, 58, 61 Friedericiella, 73, 75
Science Series 28
Ecamphicteis, 121, 125 Echinoiiiyachus, 114 Eclysippe, 123, 125 Egamella, 121, 125 Ehlersia, 81, 82 Ehlersiella, 133 Ehlersileanira, 70 Bisigella, 36 Elicodasia, 27 Enapteris, 99 Enigma, 117 Enipo, 57, 61 Enonella, 110 Enoplobranchus, 129, 131 Eone, 94 Ephesia, 98 Ephesiella, 98 Ephesiopsis, 98 Epicaste, 94 Epidiopatra, 104, 105 Epitoka, 53 Eracia, 51 Eranno, 109 Ereutho, 133 Eriographis, 140 Eriphyle, 107 Escholtzia, 99 Eteone, 47, 49 Eteonella, 51 Eteonides, 51 Eteonopsis, 113 Ethocles, 22 Euarche, 66 Eucaiphus, 147 Eucaninculatus, 103 Eucerastes, 85 Euchone, 41, 137, 138 Euchonella, 137, 138 Euclymene, 39, 40 Ettcranta, 57, 61 Eudistylia, 137, 138 Eudontosyllis, 80, 82 Euglycera, 92 Eugrymaea, 133 Eulaeospira, 148, 149, 150 Eulagisca, 58, 61 Eulalia, 48, 49 Eulalides, 51 Euleanira, 71 Eulepethidae, 8, 10, 67, 68 Eulepethus, 68 Eulepis, 68 Evimenia, 45, 51 Eumeniopsis, 45 Eumida, 48, 49 Eumidia, 51 Eamolpe, 65 Eunereis, 87, 88 Eunice, 106, 107 Eunicidae, 8, 12, 105-107, 106 Euniphysa, 106 Eunoe, 57, 61 Eunomia, 51
Fabricia, 137, 139 Fabriciella, 140 Fabriciola, 137, 139 FalTisabella, 137, 139 FaUclandiella, 17 Fauveldora, 149, 150 Fauvelia, 80, 82 Fauveliella, 73 Fauveliopsidae, 8, 13, 115, 117118 Fauveliopsis, 118 Ficopomatus, 143, 144 Filibranchus, 134 Filigrana, 147 Filograna, 142, 144 Filogranula, 142, 143, 144 Filopora, 147 Fimbriosthenelais, 69, 70 Flabellidemia, 19. 116, 117 Flabelligella, 30, 118 Flabelligera, 116, 117 Flabelligeridae, 8, 10, 13, 115, 115-117 Felmingia, 117 Flota, 118
Galathowenia, 114, 115 Galeolaria, 142, 144 Garjaiowella, 140 Gastroceratella, 65 Gastrolepldia, 59, 61 Gattiola, 85 Gattyana, 57, 62 Geminosyllis, 81. 82 Genetyllis, 48, 49 Gisela, 17 Globiodoce, 51 Glossopsis, 147 Glycera, 91,91 Glycerella, 91, 92 Glyceridae, 8, II, 14, 91-92, 92 (T13) Glycinde, 92, 93, 94 Glyphanostomum. 123, 125 Glyphohesione, 79 Gnatholycastis, 91 Gnathosyllis, 85 Goniada, 93 Goniadella, 93 Goniadidae, 8, 11, 14, 92-94. 93 (T13) Goniadides, 93 Goniadopsis, 93 Goibunovia, 140 Gorekia, 57, 62 Gossia, 85 Gravierella, 39, 40 Greeffia, 52 Giubea, 85 Grubeopolynoe, 58, 62 Grubeosyllis, 85 Gnibeulepis, 68 Grubianella, 124. 125 Giymaea, 133 Cunnarea, 118 Gwasitoa. 45 Gymnonereis, 87, 89 Gymnorhynchus, 91 Gymnosoma. 140 Gyptis. 75, 76 H Haematocleptes, 110 Halimede, 77 Haliplanella, S3 Haliplanes, S3 Halithea, 55 Halla, 111 Halodora, 52 Halogenia, 55 Halosydna, 56, 59, 62 Halosydnella, 59, 62 Halosydnoides, 65 Halosydnopsis, 59, 62 Halyplanes, 53 Hamiglycera, 92 Haplobranchus, 140 Haploscoloplos, 16 Haplosyllides. 81, 82
1976 Hapiosyllis, 81, 82 Hannopsides, 65 Harmothoe, 56, 57, 62 Haipochaeta, 79 Hartmania, 57, 62 Haswellia, 71 Hauchiella, 129, 131 Hediste, 5, 88, 89 Hedyle, 91 Hekaieiobranchus, 26 Helena, 147 Helicosiphon, 149, 150 Hemilepidia, 57, 62 Heminereis, 91 Hemipodia, 92 Hemipodus, 91,92 Hemisyllis, 85 Heptaceias, 105 Herdmanetla, 56, 62 Heimadion, 57, 62 Heimella, 120 Hermenia, 58, 62 Hermione, 55 Hermionopsis, 54, 55 Heimodice, 102 Hermonia, 55 Hermundura, 79 Hesiocaeca, 74, 76 Hesione, 75, 76 Hesionella, 74, 76 Hesionidae, 8, 12, 73-77, 74 Hesionides, 75, 76 Hesionura, 47, 49 Hesiospina, 75, 76 Hesiosyllis, 73, 76, 84 Hesperalia, 85 Hesperonoe, 57, 62 Hespeniphyllum, 47, 49 Heteraphrodita, 53, 54 Heterobranchus, 128 Heterocimis, 30 Hetenxlymene, 39, 40 Heterodisca, 151 Heteiomatdane, 41 Heteromarphysa, 107 Heteromastides, 31, 34, 36 Heteromastus, 32, 34, 36 Heteronereis, 91 HeteiDpale, 72 Heterophenacia, 133 Heterophyselia, 134 Heteropodarke, 75, 76 Heteropolynoe, 57, 62 Heterospio, 26, 27 Heterospionidae, 8, 14, 27-28, 29 Heterosyllis, 85. Heteroterebella, 134 Hipponoa, 100, 102 Histriobdella, 114 Histriobdellidae, 8, 12, 113, 113114 Histriodrilus, 114 Hololepida, 59, 62 Hololepidella, 58, 62 Horstileanira, 69, 70 Hyalinoecia, 104, 105 Hyalopomatopsis, 147 Hyalopomatus, 143, 144
755
Labotas, 17 Labranda, 30 Labrorostratus, 110 Lacharis, 94 Lacydes, 17 Lacydonia, 94 Lacydoniidae, 8, 12, 94, 94 Ladice, 43 Laenilla, 65 Laeonereis, 87, 89 Laeospira, 151 Laetmatonice, 55 Laetmonioe, 54 Lagis, 120 Lagisca, 57, 63 Lais, 111 Lalage, 85 Lamellisyllis, 80, 83 Lanassa, 129, 131 Langerhansia, 65, 85 Lanice, 130, 131 Lanicides, 130, 131 Laomedora, 85 Laonice, 23, 24 Laonome, 137, 139 Laonomedes, 140 Laphania, 129, 131 Laphaniella, 134 Lapithas, 85 Laranda, 111 Laiymna, 111 Leaena, 129, 131 Leanira, 69, 70 Ledon, 31 Leiobranchus, 140 Lîocapitella, 33, 34, 36 Leiocapitellides, 32, 34, 36 Leiocephalus, 41 Leiochone, 41 Leiochrides, 32, 34, 36 Leiochrus, 32, 34, 36 Leipoceras, 26 Lenora, 103 Leocrates, 75, 76 Leocratides, 75, 76 Leodice, 107 Leodora, 148, 149, 150 Leoimates, 87, 89 Leonnatus, 94 Leontis, 91 Lepadorhynchus, 51 Lepidametria, 65 Lepidasthenia, 59, 63 Lepidastheniella, 59, 63 Lepidia, 66 Lepidombria, 58, 63 Lepidogyra, 58, 63 Lepidonereis, 91 Lepidonotus, 56, 58, 63 Lepidopleunis, 71 Lepraea, 134 Leptochaetoptenu, 29 Leptochone, 140 Leptoecia, 105 Leptoneieis, 87, 89 Letmonicella, 55 Leucariste, 134 Leucia, 57, 63
Hyboscolex, 44 Hydroides, 143, 144, 147 Hydrophane, 53 Hypania, 124, 125 Hypaniola, 124, 125 Hypephesia, 98 Hypereteone, 51 Hyperhalosydna, 59, 62 Hypociirus, 51 Hypoeulalia, 51 Hypsicomatides, 140 Hypsicomatopsis, 140 Hypsicomus, 137, 139 I Ichthyotomidae, 8, 11, 113, 114 Ichthyotomus, 114 Idalia, 134 Idanthyrsus, 118 Inermonephtys, 97 Ilyphagus, 116, 117 Intoshella, 57, 63 loda, 85 lospilidae, 8, 14, 95, 95-96 lospilopsis, 95, 96 lospilus, 95, 96 Iphianissa, 41 Iphinereis, 91 Iphione, 56, 63 Iphionella, 65 Iphitime, 109 Iphitimidae, 8, 11, 108, 109-110 Irana, 121, 126 Irmula, 80, 83, 84 IsociiTus, 39, 40 Isolda, 121, 126, 127 Isoma^tus, 36 Isosyllis, 85
Janita, 143, 144 Janua, 149, 150 Jasmineira, 137, 139 Johnstonella, 99 Johnstonia, 39, 40 Josephella, 141, 144
Kainonereis, 87, 89 Kalaminochaeta, 27 Kalaminochaetidae, 27 Kalununaria, 27 Kebuita, 44 Kefersteinia, 75, 76 Kermadecella, 57, 63 Kesun, 42, 43 Kinbergella, 26 Kinbergia, 51 Kinberginereis, 87, 89 Krohnia, 51, 52 Kuwaiia, 107, 109 Kynephorus, 79
Labiaria, 121 Labidognathus, HI
184
NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY Mandane, 26 Mandrocles, 41 Mania, 77 Marenzelleria, 23, 24 Marifugia, 142, 145 Marphysa, 106, 107 Marsipospira, 151 Mastigethus, 53 Mastigonereis, 91 Mastobranchus, 32, 34, 36 Mauita, 52 Maupasia, 53 Mayella, 69, 70 Mayeria, 107 Mediomastus, 32, 34, 36 Megachone, 140 Megalomma, 137, 139 Meganereis, 91 Meganerilla, 152, 154 Meiodotviltea, 112 Melaenis, 57, 63 Melinantipoda, 121, 126 Melinella, 130, 131 Melinna, 121, 126 Melinnampharete, 123, 126 Melinnata, 123, 126 Melinnexis, 121, 126 Melinnides, 128 Melinnoides, 123, 126 Melinnopsides, 121, 126 Melinnopsis, 121, 126, 128 Membranopsis, 145, 147 Mera, 151 Mercierella, 143, 145 Mercierellopsis, 143, 145 Mesochaetopterus, 28 Mesocirrineris, 30 Mesoeulalia, 51 Mesomystides, 51 Mesonerilla, 152, 154 Mesospio, 23, 24 Mesotrocha, 29 Metachone, 140 Metalaeospira, 149, 150 Metalaonome, 140 Metamphinome, 103 Metavermilia, 142, 145 Mexamage, 124, 126 Mexieulepis, 68 Microclynaene, 39, 40 Micromaldane, 38, 40 Micronephthys, 97 Micronereides, 87, 89 Micronereis, 87, 89 Microphthalmus, 75, 76 Micropodarke, 75, 76 Microrbinia, 15, 16 Microsamytha, 128 Microserpula, 147 Microspio, 23, 24 Microsyllis, 85 MUnesia, 55 Minuspio, 23, 24 Mitrana, US Monocerina, 85 Monorchos, 118, 119, 120 Monostylos, 20 Monroika, 137, 139, 140 Monticellina, 30 Morants, 22, 24 Moyanus, 121, 126 Mugga, 123, 126 Muggoides, 123, 126 Myriana, 51 Myrianida, 79, 83 Myriacyclum, 51 Myriochele, 115 Myrioglobula, 115 Myriowenia, 114, 115 Mysta, 47, 49 Mystides, 47, 49 Myxicola, 136, 139 N
Science Series 28
Leucodora, 26 Levidorum, 97, 98 Levinsenia, 18 Linophenis, 100, 102 Liocapa, 52 Lipephila, 91 Lipobranchius, 43, 44 Lirione, 103 Lithognatha, 107 Litocorsa, 78 Loandalia, 78 Loimia, 130, 131 Longosoma, 28 Lopadorhynchidae, 8, 12, 46, 5253 Lopadorhynchus, 53 Lophocephalus, 117 Lophonota, 103 Lophosyllis, 85 Lucopia, 58, 63 Lugia, 47, 49 Lumara, 134 Lumbriclymene, 38, 40 Lumbriclymenella, 38, 40 Lumbricomastus, 33, 34, 36 Lumbriconais, 36 Lumbriconereis, 109 Lumbrineridae, 8, 13, 107-109, 108 Lumbrinendes, 109 Lumbrineriopsis, 109 Lumbrineris, 107, 109 Lycaretus, 103 Lycastella, 91 Lycastis, 85, 91 Lycastoides, 87, 89, 91 Lycastopsis, 87, 89 Lycidice, 107 Lycoris, 91 Lygdamis, 118 Lysarete, 111 Lysaretidae, 8, 11, 108, 111 Lysidice, 106 Lysilla, 129, 131 Lysippe, 124, 126 Lysippides, 124, 126
M
Macduffia, 107 Macellicephala, 56, 63 Macellicephaloides, 56, 63 Macelloides, 56, 63 Maclovia, 111 Macrochaeta, 30, 31 Macroclymene, 39, 40 Macroclymenella, 39, 40 Macrophyllum, 51 Maea, 26 Mageiona, 26 (TIO) Magelonidae, 8, 14, 26, 26 Malacoceros, 23, 24 Maldane, 39, 40 Maldanella, 39, 40 Maldanidae, 8, 13, 37-41,38 Maldanopsis, 41 Malmgrenia, 59, 63 Malmgreniella, 59, 63 Manayunkia, 137, 139, 140
Naiades, 51, 52 Naidonereis, 17 Nainereis, 15, 16 Nais, 43 Naraalycastis, 87, 89 Namanereis, 87, 89 Naneva, 130, 131 Naraganseta, 30 Naumachius, 91 Nauphanta, 52, 107 Nausicaa, 107 Neanthes, 86, 88, 89 Neco, 41 Nectochaeta, 65 Nectoneanthes, 88, 89 Nectonereis, 91 Nematonereis, 106, 107 Nemidia, 57, 64 Neoamphitrite, 129, 131 Neodexiospira, 151 Neoheteromastus, 32, 34, 36 Neohololepidella, 58, 64 Neoleanira, 69, 70 Neoleprea, 129, 132 Neolipobranchius, 43, 44 Neomediomastus, 32, 34, 36 Neomeris, 43 Neomicrorbis, 148, 149, 151 Neonotomastus, 32, 34, 36 Neopaiwa, 123, 126 Neopanthalis, 66 Neopodarke, 75, 76 Neopomatus, 143, 145 Neopygospio, 26 Neosabellides, 123, 127 Neosamytha, 123, 127 Neottis, 134 Neovermilia, 143, 145 Nephtyidae, 8, 12, 95, 96, 96-97 (TI3) Nephtys, 97 Nereidae, 8, 12, 85-91, 86 Nereidice, 107 Nereidonta, 107 Neieilepas, 91 Nereimyra, 75, 76 Nereiphylla, 48, 49 Nereis, 86, 88, 90 Nereisyllis, 85 Nereitube, 105 Nerilla, 152. 154
W6
Nerillidae. 8, 12, 152-154, 253 Nerllidum, 154 Nerillidopsis, 154 Nerine, 26 Nerinides, 26 Nerinopsis, 26 Nevaya, 27 Nicolea, 130, 132 Nicomache, 38, 40 Nicomachella, 41 Nicomedes, 91 Nicon, 87, 90 Nicotia, 85 Ninoe, 107, 109 Nipponophyllum, 47, 50 Nitetis, 43 Norepa, 65 Northia, 105 Nossis, 91 Notalia, 48, 50 Notaulax, 140 Nothis, 51 Nothria, 105 Notocirrus, 110, 111 Notodasus, 32, 34, 36 Notomastus, 32, 34, 36 Noiophyllum, 47, 50 Notoproctus, 38, 41 Notopsilus, 110, 111 Notopygos, 100, 102 Novaquesta, 19 Novobranchus, 134 Nuchubranchiata, 100 Nudisyllis, 80, 83 Nychia, 65 O Octobranchus, 134 Odontosyllis, 81, 83 Odysseus, 134 Oenone, 111 Oeorpata, 127 Olga, 142, 145 Olgahannania, 143, 145 Oligobranchus, 45 Otigognathus, 110 Oligolepis, 65 Omaria, 43 Omphalopoma, 143, 145 Omphalopomopsis, 142, 145 Oncoscolex, 45 Oniscosoma, 104 Onuphidae, 8, 10, 12, 104, 104-105 Onuphis, 105 Oophylax, 85 Ophelia, 42, 43 Opheliidae, 8, 14, 41^3, 42 Ophelina, 42, 43 Ophiodromus, 75, 76, 77 Ophioglycera, 93, 94 Ophiuricola, 107, 109 Ophryotrocha, 112 Opisthodonta, 80, 83 Opisthopista, 129, 132 Opisthosyllis, 81, 83 Ops, 115 Orbinia, 16
1&5
Parapolynoe, 65 Paraprionospio, 23, 24 Parapterosyllis, 81, 83 Parasabella, 140 Parascierocheus, 44 Paraserpula, 143, 145 Parasphaerosyllis, 81, 83 Parasitosyllis, 85 Parasyllidea, 75, 77 Parathelepus, 129, 132 Paiatyposyllis, 81, 83 Parautolytus, 80, 83 Paraveimilia, 147 Paraxionice, 130, 132 Paraxiothea, 41 Parergodrilidae, 8, 13, 19,20 Parergodrilus, 20 Pareulepis, 68 Pareupholoe, 69, 70 Pareurythoe, 100, 102 Parexogone, 85 Parheteromastides, 32, 35, 36 Parheteromastus, 32, 35, 36 Parhypania, 124, 127 Patmensis, 65 Paronuphis, 105 Parophryotrocha, 112 Parthenope, 20 Paumotella, 142, 145 Pectinaria, 120 Pectinariidae, 8, 10, 119, 120-121 Peduiosoma, 53 Peisidice, 68 Peisidicidae, 66 Pelagobia, 52, 53 Pellucidaria, 97 Pelogenia, 71 Pentacirrus, 30 Peresiella, 31, 35, 36 Perialla, 26 Periboea, 73, 77, 85 Perinereis, 88, 90 Perolepis, 59, 64 Petaloclymene, 41 Petaloproctus, 39, 41 Petitia, 80, 83 Petta, 120, 121 Phaetusa, 91 Phalacrophorus, 95, 96 Phalacrostemma, 118, 119 Pharyngeovalvata, 80, 83 Phenacia, 134 Pherea, 134 Pherecardia, 102, 103 Pherecardites, 102, 103 Pherasa, 116, 117 Phisidia, 129, 132 Pholpe, 69, 70 Pholoides, 66 Pholoididae, 8, 10, 66, 67 Photocharis, 85 Phragmatopoma, 118, 119 Phronia, 79 Phyllampharete, 123, 127 Phyllamphicteis, 124, 127 Phyllochaetopterus, 28 Phyllocomus, 124, 127 Phyllodoce, 48, 50, 51
Orbiniella, 15, 16 Orbiniidae, 8, 13. 14-17, IS Oria, 140 Oriades, 140 Oridia, 140 Oriopsis, 137, 139 Orseis, 73, 76 Orthodromus, 77 Otanes, 134 Othonia, 140 Otopsis, 78 Owenia, 114, 115 Oweniidae, 8, 11, 14, 114-115, 115 Oxydromus, 77
Pabits, 123, 127 Paedophylax, 85 Pagenstecheria, 85 Paiwa, 124, 127 Paleanotus, 72 Pallasia, 120 Pallasina, 120 Pallonia, 134 Palmyra, 72 Palmy reuphrosyne, 103 Palmyridae, 8, 10, 71, 71,72 Palola, 106, 107 Palpiglossus, 107 Panthalis, 66 Panousea, 138, 139 Pantoithrix, 116, 117 Papillaria, 26 Parachloeia, 100, 102 Paracbonia, 140 Paradexiospira, 149, 151 Paradiopatra, 105 Paradoneis, 18 Paradyte, 57, 64 Paraeulalia, 51 Paiahalosydna, 58, 64 Parahesione, 75, 77 Parahololepidella, 58, 64 Paraiospilus, 95, 96 Paralacydonia, 94 Paralaeospira, 149, 151 Paralanice, 130, 132 Paraleiocapitella, 32, 35, 36 Paralepidonotus, 59, 64 Paralycastis, 91 Paramage, 124, 127 Paramarphysa, 106, 107 Paramphicteis, 124, 127 Paramphinome, tOO, 102 Paranaitis, 48, 50 Parandalia, 78, 79 Paranereis, 91 Paranerilla, 154 Paranerine, 26 Paranorthia, 104, 105 Paranychia, 65 Paraonella, 18 Paraonidae, 8, 13, 14, 15, 17-lS Paraonides, 18 Paraonis, 18 Parapholoe, 68 Parapionosyllis, 80, 83
]86
NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY Pomatoceros, 142, 146 Pomatoleios, 142, 146 Pomatostegus, 142, 146 Pontodora, 53 Pontodoridae, 8, 12, 46, 53 Pontogenia, 54 Poitoa, 51 Portelia, 97 Potamethus, 138, 139 Potamilla, 138. 139, 140 Potamis, 140 Potaspina, 138, 139 Pottsiscalisetosus, 58, 64 Praegeria, 73 Praxilla, 41 PraxUlella, 39, 41 Praxillura, 38, 41 Praxithea, 91 Prionognathus, 113 Prionospio, 22, 23, 24 Proboscidea, 92 Proceraea, 79, 83 Procerastea, 79, 84 Prochaetoparia, 48, 50 Proclea, 129, 132 Proclymene, 39, 41 Procome, 85 Prodexiospiia, 149, 151 Progoniada, 92, 94 Promaldane, 41 Promenia, 30 Prophyllodoce, 48, 50 Proplacostegus, 147 I^scalibregma, 43, 44 Proscoloplos, 15, 16 EVospio, 26 Protis, 141, 146 Protoaricia, 15, 16 Protoariciella, 15, 17 Protocapitella, 37 Protocarobia, 51 Protodorviliea, 112 Protcxlrilidae, 8, 9, 12, 153, 155 Protodriloides, 155 E>rotodrilus, 155 Ftetogrubea, 85 Protolaeospira, 149, 151 Protoleodora, 149, 151 Ehotolycoris, 91 Protomastobranchus, 33, 35, 36 Protomystides, 48, 50 Protopolydora, 26 Protothelepus, 134 Protula, 146 Protulides, 140 Psammocollus, 115 Psammodrilidae, 8, 13, 20, 20-21 Psamraodriloides, 21 Psammodrilus, 21 Psammolyce, 69, 70 Psammoriedlia, 154 Psectra, 72 Pseudampharete, 128, 132, 133 Pseudeulalia, 48, 50 Pseudeupanthalis, 66, 71 Pseudeurythoe, 103 Pseudexogone, 80, 84 Pseudobranchiomma, 137, 140
Science Series 28
Phyllodocidae, 8, 12, 45-51, 46 (T-13) Phyllohattmania, 57, 64 Phyllonereis, 91 Phyllosheila, 57, 64 PhyllosylHs, 79, 83 Phylo, 16 Physalidonotus, 65 Phyzelia, 134 PUargiidae, 8, 12, 74, 77-79 Pilargis, 78 Pilearia, 27 Peolaria, 149, 151 PUlaiospira, 148, 149, 151 Pionosyllis, 80, 83 Pirakia, 48, 50 Piratesa, 147 Piromis, 116, 117 Pisenoe, 91 Pisione, 73 Pisionella, 73 Pisionidae, 8, 10, 12, 72, 72-73 E^sionidens, 73 Pisionura, 72 Pista, 130, 132, (TO) Pixellia, 152 Placostegus, 142, 145 Placostegopsis, 147 Plakosyllis, 80, 83 Platyneteis, 86, 88, 90 Platysyllis, 85 Pleione, 103 Plioceras, 111 Plotobia, 100 Plotohelmis, 51, 52 Plotolepis, 65 Podarke, 75, 77 Podarkeopsis, 75, 77 Podarmus, 58, 64 Podioceros, 147 Podonereis, 85, 91 Poecilochaetidae, 8, 14, 27, 27 Poecilochaetus, 27 Poeobiidae, 8,9, 115, 117 Poeobius, 117 PoUicita, 98 Polybostrichus, 85 Polybranchia, 22, 24 Polycimis, 129, 132 Polydora, 22, 24, 25 Polydorella, 22, 24 Polyeunoa, 58, 64 Polygoidiidae. 8, 153, 154-155 Polygordius, 154, 155 Polylepis, 71 Polymastus, 85 Polymnia, 134 Polymniella, 129, 132 Polynice, 85 Polynoe, 57, 64 Polynoella, 56, 64 Polynoidae, 8, 10, 55, 55-65 Poiyodontes, 66 Poiyodontidae, 8, 10, 55, 65-66 Polyophthalmus, 42, 43 Polyphragma, 147 Poiyphysia, 43, 44 Pomatoceropsis, 147
Pseudocapitella, 33, 35, 36 Pseudochitinopoma, 143, 146 Pseudocirratulus, 29, 30 Pseudoclymene, 39, 41 Pseudofabricia, 137, 140 Pseudohalosydna, 59, 64 Pseudolacydonia, 94 Pseudoleiocapitella, 32, 35, 36 Pseudoleucodore, 26 Pseudomalacoceros, 23, 24 Pseudonereis, 88, 90 Pseudonerine, 26 Pseudonotophyllum, 51 Pseudopalolo, 107 Pseudopolydora, 22, 25 Pseudopolynoe, 58, 65 Pseudopomatoceros, 142, 146 Pseudopotamilla, 140, 141 Pseudosabellides, 128 Pseudoscalibregma, 44 Pseudoserpula, 141, 146 Pseudosyllides, 81, 84, 85 Pseudosyllidia, 77 Pseudothelepus, 133 Pseudovefmilia, 143, 146 Psymobranchus, 147 Pterampharete, 123, 127 Pterautolytus, 85 Pteriptyches, 26 PterociiTus, 48, 50 Pterolysippe. 124, 127 Pteroscolex, 37 Pterosyllis, 85 Pterothrix, 111 Pulliella, 32, 35, 36 Puparia, 37 Pycnoderma, 117 I^gophelia, 43 Pygophyllum, 26 F^gospio, 23, 25
Questa, 19 Questidae, 8, 13, IS, 18-19 Quetieria, 65
Ramex, 130, 132 Ranzania, 29 Ranzanides, 29 Raphidrilus, 20 RariciiTUS, 29, 30 Rashgua, 32, 35, 36 Reibischia, 53 Restio, 66 Reteterebella, 130, 132 Rhamphobrachium, 104, 105 Rhodine, 38, 41 Rhodopsis, 142, 146 Rhopalosyllis, 80, 84 Rhynchobolus, 92 Rhynchonerella, 51, 52 Rhynchospio, 23, 25 Rhynophylla, 26 Robertianella, 56, 65 Romanchella, 149, 151 Rostrada, 103
1976 RuUierinereis, 87, 90 Rytocephalus, 128
187 Subprotula, 141, 147 Syllia, 85 Syllidae, 8, 12, 74, 79-85 Syllides, 80, 84 Syllidia, 75, 77 Sylline, 85 Syllis, 81, 84 Synelmis, 78 Synsyllis, 80, 84
Sabaco, 41 Sabella, 137, 140 Sabellaria, 118, 119, 120 Sabellariidae, 8, 10, 118-120, 119 Sabellastarte, 137, 140 Sabellidae, 8, 11, 135-141, 136 Sabellides, 123, 127 Sabellina, 140 Sabellonga, 141 Sabellongidae, 8, 13, 136, 141 Sabidius, IS Sabina, 140 Saccocirridae, 8, 153, 155 Saccocirrus, 155 Sacconereis, 85 Sagiteila, 99 Salmacina, 141, 146 Salmacinopsis, 141, 146 Salvatoria, 85 Samytha, 123, 127 Samythella, 123, 127 Samythopsis, 123, 128 Sandanis, 36 Sangiria, 100, 103 Saphobranchia, 117 Scalibregma, 44 Scaiibregmella, 44, 45 Scalibregmidae, 8, 14, 42, 43^5 Scalibregmides, 44, 45 Scalis, 121 Scalisetosus, 57, 65 Scalispinigera, 43, 44, 45 Schistocomus, 124, 128 Schistomeringos, 112, 113 Schizobranchia, 137, 140 Schizocraspedon, 142, 146 Schmardanella, 134 Schmardia, 85 Schmardiella, 77 Schreiberius, 114 Schroederella, 15, 17 Scione, 134 Scionelia, 130, 132 Scionides, 130, 132 Scionopsis, 134 Sclerobregma, 44, 45 Sclerocheilus, 44, 45 Sclerostyta, 142, 143, 146 Scolecolepides, 23, 25 Scolecolepis, 26 Scoleiepis, 23, 25 Scoletoma, 109 Scoloplella, 15, 17 Scoloplos (Leodamas) 16, 17 Scoloplos (Scoloplos) 16, 17 Scyphoproctus, 32, 35, 36 Semiodera, 117 Serpula, 143, 147 Serpulidae, 8, 10. 11, 141-147, 148 Sheila, 58, 65 Sigalion, 69, 70 Sigalionidae, 8. 10, 68-71, 69 Sigambra, 78
Sige, 48, 50 Siliquaria, 147 Simplicaria, 148, 149, 151 Sinistrella, 152 Siphonostoma, 117 Siphostoma, 117 Sttophaga, 141 Skardaria, 22 Solomononereis, 87, 90 Solowetia, 134 Sonatsa, 39, 41 Sosane, 124, 128 Sosanella, 123, 128 Sosanides, 124, 128 Sosanopsis, 124, 128 Spermosyllis, 80, 84 Sphaerephesia, 97, 98 Sphaerodoce, 48, 50 Sphaerodoridae. 8, 12, 95, 96, 9798 Sphaerodoridium, 98 Sphaerodoropsis, 98 Sphaerodorum, 98 Sphaeropomalus, 143, 147 Sphaerosyllis, 80, 84, 97 Spinosphaera, 129, 132 Spinther, 100, 104 Spintheridae, 8, 9, 100, 101, 104 Spio, 23, 25 Spiochaetopterus, 28 Spione, 26 Spionereis, 26 Spionidae, 8, 14, 21, 22-26 Spionides, 26 Spiophanes, 22, 25 Spiramella, 147 Spirobranchus, 142, 147 Spirodiscus, 142, 147 Spirographis, 137, 140 Spirorbella, 148, 149, 151 Spirorbidae, 8, 11, 147-152, 148 Spirorbides, 152 Spirorbis, 149, 151 Spirovenna, 129, 133 Staurocephalus, 113 Stauroceps, 113 Stauronereis, 113 Steggoa, 48, 50 Steninonereis, 87, 90 Stephania, 77 Stephanosyllis, 85 Stemaspidae, 8, 10, 113, 114 Stemaspis, 114 Sthenelais, 70 Sthenelanella, 69, 70 Sthenolepis, 70 Stoa, 152 Strategis, 103 Stratiodrilus, 114 Stratonice, 91 Sireblosoma, 129, 133 Streblospio, 23, 25 Streptosyllis, 80, 84 Struwela, 74, 77 Stschapoveila, 129, 133 Stygocapitella, 20 Stylarioides, 117 Subadyte, 57, 65
Tachy try pane, 42, 43 Tainokia, 111 Talehsapia, 78 Taphus, 72 Tauberia, 18 Tecturella, 117 Telake, 92 Telamone, 77 Teiepsavus, 29 Telethusae, 37 Telolepidasthenia, 59, 65 Telonereis, 113 Telothelepus, 129, 133 Temporaria, 142, 147 Tenonia, 57, 65 Terebella, 129, 133 Teiebellanice, 130, 133 Teiebellidae, 8, 11, 122, 128-134 Terebellides, 134 Terebellobranchia, 129, 133 Terpsichore, 43 Tetraglene, 85 Tetratrocha, 91 Tetreres, 120 Thalassema, 114 Thalassochaetus, 154 Thalenessa, 69, 70 Tharyx, 29, 30 Thaumastoma, 27 Thelepella, 134 Thelephusa, 134 Thelepides, 130, 133, 134 Thelepodopsis, 134 Thelepus, 129, 133 Theodisca, 17 Theostoma, 17 Therochaeta, 116, 117 Therochaetella, 116, 117 Thesmia, 103 Thetisella, 103 Thoe, 85 Thoosa, 91 Thormora, 58, 65 Thylaciphorus, 85 Thysanoplea, 98 Tibiana, 107 TimaiBte, 29, 30 Tomopteridae, 8, II, 95, 96, 98-99 Tomopteris, 99 Torquea, 134 Torrea, 51, 52 Trachelophyllum, 51 Trapodia, 105 Travisia, 42, 43 Travisiopsis, 99 Tricertia, 53, 54
188
NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY U Umbellisyllis, 79, 84 Unanereis, 87, 90 Uncinerels, 91 Uncinia, 26 Unciniseta, 109 Uncinochaeta, 134 Uncopolynoe, 56, 65 Unobranchus, 134, 135 Urosiphon, 43 V Valla, 36 Vanadis, 51, 52 Veleda, 103 Venadis, 17 Venusia, 134 Vennilia, 147 Venniliopsis, 143, 147 Virchowia, 85 Vitiazia, 48, 50 Xenosyilides, 85 Xenosyllis, 81, 84 ^ Zeppelinia, 20 Zopyrus, 147 Zorus, 117 Zothea, 103 Zygolobus, 109 Zygophyllus, 109
Science Series 28
Trichobranchidae, 8, 11, 122, 134135 Trichobranchus, 134, 135 Trichosobranchella, 140, 141 Trichosyllis, 85 Tricoelia, 29 Tricosmoehaeta, 65 Trilobodrilus, 152 Tripolydora, 22, 25 Trochochaeta, 26 Trochochaetidae, 8, 14, 21, 26-27 Troglochaetus, 154 Trophonia, 117 Trophoniella, 116, 117 Ttypanosyllis, 79, 81, 84 Tylonereis, 87, 90 Tylorrhynchus, 87, 90 Typhlonereis, 91 Typhloseolecidae, 8, 14, 95, 99, 99-100 Typhloscolex, 99 Typosyllis, 81, 84 Tyrrhena, 77
Vitiaziphyllum, 48, 51 W Wartelia, 134 Watelio, 51 52 Webena, 59, 65 ^^T'Tî' o Weddellia, 123, 12S Wesenbergia 75 77 Wleysthenelais, 69, 70 ^

The Polychaete Worms. Definitions and Keys To The Order, Families and Genera

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THE POLYCHAETE WORMS

DEFINITIONS AND KEYS TO THE ORDERS, FAMILIES AND GENERA

Science Series 28 February 3, 1977