5. »3 : : i CI y*l NOAA TR NMFS CIRC-374 A UNITED STATES DEPARTMENT OF COMMERCE PUBLICATION j*» T ' \ CT NOAA Technical Report NMFS CIRC-374 U.S. DEPARTMENT OF COMMERCE National Oceanic and Atmospheric Administration National Marine Fisheries Service Marine Flora and Fauna of the Northeastern United States, Annelida: Oligochaeta DAVID G. COOK and RALPH 0. BRINKHURST SEATTLE, WA May 1973 NOAA TECHNICAL REPORTS National Marine Fisheries Service, Circulars The major responsibilities of the National Marine Fisheries Service (NMFS) are to monitor and assess the abundance and geographic distribution of fishery resources, to understand and predict fluctuations in the quan- tity and distribution of these resources, and to establish levels for optimum use of the resources. NMFS is also charged with the development and implementation of policies for managing national fishing grounds, develop- ment and enforcement of domestic fisheries regulations, surveillance of foreign fishing off United States coastal waters, and the development and enforcement of international fishery agreements and policies. NMFS also assists the fishing industry through marketing service and economic analysis programs, and mortgage insurance and vessel construction subsidies. It collects, analyses, and publishes statistics on various phases of the industry. The NOAA Technical Report NMFS CIRC series continues a series that has been in existence since 1941. The Circulars are technical publications of general interest intended to aid conservation and management. Publica- tions that review in considerable detail and at a high technical level certain broad areas of research appear in this series. Technical papers originating in economics studies and from management investigations appear in the Circular series. NOAA Technical Reports NMFS CIRC are available free in limited numbers to governmental agencies, both Federal and State. They are also available in exchange for other scientific and technical publications in the ma- rine sciences. Individual copies may be obtained (unless otherwise noted) from NOAA Publications Section, Rock- ville, Md. 20852. Recent Circulars are: 315. Synopsis of biological data on the chum salmon, Oncorhynchus keta (Walbaum) 1792. By Rich- ard G. Bakkala. March 1970, iii + 89 pp., 15 figs., 51 tables. 319. Bureau of Commercial Fisheries Great Lakes Fishery Laboratory, Ann Arbor, Michigan. By Bureau of Commercial Fisheries. March 1970, 8 pp., 7 figs. 330. EASTROPAC Atlas: Vols. 4, 2. Catalog No. 1 49.4:330/ (vol.) 11 vols. ($4.75 each). Avail- able from the Superintendent of Documents, Washington, D.C. 20402. 331. Guidelines for the processing of hot-smoked chub. By H. L. Seagran, J. T. Graikoski, and J. A. Emerson. January 1970, iv -f 23 pp., 8 figs., 2 tables. 332. Pacific hake. (12 articles by 20 authors.) 1970, iii + 152 pp., 72 figs., 47 tables. March 333. Recommended practices for vessel sanitation and fish handling. By Edgar W. Bowman and Alfred Larsen. March 1970, iv + 27 pp., 6 figs. 335. Progress report of the Bureau of Commercial Fisheries Center for Estuarine and Menhaden Research, Pesticide Field Station, Gulf Breeze, Fla., fiscal year 1969. By the Laboratory staff. August 1970, iii + 33 pp., 29 figs., 12 tables. 336. The northern fur seal. By Ralph C. Baker, Ford Wilke, and C. Howard Baltzo. April 1970, iii + 19 pp., 13 figs. 337. Program of Division of Economic Research, Bureau of Commerecial Fisheries, fiscal year 1969. By Division of Economic Research. April 1970, iii + 29 pp., 12 figs., 7 tables. 338. Bureau of Commercial Fisheries Biological Lab- oratory, Auke Bay, Alaska. By Bureau of Com- mercial Fisheries. June 1970, 8 pp., 6 figs. 339. Salmon research at Ice Harbor Dam. By Wesley J. Ebel. April 1970, 6 pp., 4 figs. 340. Bureau of Commercial Fisheries Technological Laboratory, Gloucester, Massachusetts. By Bu- reau of Commercial Fisheries. June 1970, 8 pp., 8 figs. 341. Report of the Bureau of Commercial Fisheries Biological Laboratory, Beaufort, N.C., for the fiscal year ending June 30, 1968. By the Lab- oratory staff. August 1970, iii -f- 24 pp., 11 figs., 16 tables. 342. Report of the Bureau of Commercial Fisheries Biological Laboratory, St. Petersburg Beach, Florida, fiscal year 1969. By the Laboratory staff. August 1970, iii + 22 pp.,*20 figs., 8 tables. 343. Report of the Bureau of Commercial Fisheries Biological Laboratory, Galveston, Texas, fiscal year 1969. By the Laboratory staff. August 1970, iii + 39 pp., 28 figs., 9 tables. 344. Bureau of Commercial Fisheries Tropical Atlan- tic Biological Laboratory progress in research 1965-69, Miami, Florida. By Ann Weeks. Oc- tober 1970, iv + 65 pp., 53 figs. 346. Sportsman's guide to handling, smoking, and pre- serving Great Lakes coho salmon. By Shearon Dudley, J. T. Graikoski, H. L. Seagran, and Paul M. Earl. September 1970, iii + 28 pp., 15 figs. 347. Synopsis of biological data on Pacific ocean perch, Sebastodes alutus. By Richard L. Major and Herbert H. Shippen. December 1970, iii + 38 pp., 31 figs., 11 tables. Continued on inside back cover. WMOspii r MENl Of U.S. DEPARTMENT OF COMMERCE Frederick B. Dent, Secretary NATIONAL OCEANIC AND ATMOSPHERIC ADMINISTRATION Robert M. White, Administrator NATIONAL MARINE FISHERIES SERVICE NOAA Technical Report NMFS CIRC-374 Marine Flora and Fauna of the Northeastern United States. Annelida: Oligochaeta DAVID G. COOK and RALPH 0. BRINKHURST i a S i SEATTLE, WA May 1973 For sale by the Superintendent of Documents, U.S. Government Printing Office Washington, D.C., 20402 - Price 3 5 cents FOREWORD This issue of the "Circulars" is part of a subseries entitled "Marine Flora and Fauna of the Northeastern United States." This subseries will consist of original, illustrated, modern manuals on the identification, classification, and general biology of the estuarine and coastal marine plants and animals of the Northeastern United States. Manuals will be published at irregular intervals on as many taxa of the region as there are specialists willing to collaborate in their preparation. The manuals are an outgrowth of the widely used "Keys to Marine Invertebrates of the Woods Hole Region," edited by R. I. Smith, published in 1964, and produced under the auspices of the Systematics-Ecology Program, Marine Biological Laboratory, Woods Hole, Mass. Instead of revising the "Woods Hole Keys," the staff of the Systematics- Ecology Program decided to expand the geographic coverage and bathymetric range and produce the keys in an entirely new set of expanded publications. The "Marine Flora and Fauna of the Northeastern United States" is being prepared in collaboration with systematic specialists in the United States and abroad. Each man- ual will be based primarily on recent and ongoing revisionary systematic research and a fresh examination of the plants and animals. Each major taxon, treated in a separate manual, will include an introduction, illustrated glossary, uniform originally illustrated keys, annotated check list with information when available on distribution, habitat, life history, and related biology, references to the major literature of the group, and a system- atic index. These manuals are intended for use by biology students, biologists, biological ocean- ographers, informed laymen, and others wishing to identify coastal organisms for this region. In many instances the manuals will serve as a guide to additional information about the species or the group. Geographic coverage of the "Marine Flora and Fauna of the Northeastern United States" is planned to include organisms from the headwaters of estuaries seaward to approximately the 200-m depth on the continental shelf from Maine to Virginia, but may vary somewhat with each major taxon and the interests of collaborators. When- ever possible representative specimens dealt with in the manuals will be deposited in reference collections of the Gray Museum, Marine Biological Laboratory, and other universities and research laboratories in the region. After a sufficient number of manuals of related taxonomic groups have been pub- lished, the manuals will be revised, grouped, and issued as special volumes. These vol- umes will thus consist of compilations of individual manuals within phyla such as the Coelenterata, Arthropoda, and Mollusca, or of groups of phyla. CONTENTS Page Introduction 1 Definitions and diagnostic characters 1 Ecology 4 Collecting methods 5 Examination procedure 5 Key to the marine Oligochaeta of the east coast of North America 6 Annotated systematic list 18 Selected bibliography 20 Index to scientific names 22 Acknowledgments 23 Coordinator's comments 23 The National Marine Fisheries Service (NMFS) does not approve, rec- ommend or endorse any proprietary product or proprietary material mentioned in this publication. No reference shall be made to NMFS, or to this publication furnished by NMFS, in any advertising or sales pro- motion which would indicate or imply that NMFS approves, recommends or endorses any proprietary product or proprietary material mentioned herein, or which has as its purpose an intent to cause directly or indirectly the advertised product to be used or purchased because of this NMFS publication. Ill MARINE FLORA AND FAUNA OF THE NORTHEASTERN UNITED STATES Annelida: Oligochaeta DAVID G. COOK 1 and RALPH O. BRINKHURST 2 ABSTRACT The manual includes an introduction on the general biology, an illustrated key, an an- notated systematic list, a selected bibliography, and an index to the marine Oligochaeta of the East Coast of North America. The Families Naididae, Tubificidae, Enchytraeidae, and Megascolecidae are treated. INTRODUCTION The Oligochaeta, a group of predominantly terrestrial and freshwater annelids, includes a number of species which have adapted well to the marine benthic environment. Despite the fact that these organisms may occur in great num- bers in the sea (densities of up to a million per m 2 have been reported) the information on their ecology and taxonomy is comparatively sparse. The available data do suggest, however, that the intertidal zone, the subtidal zone, the con- tinental shelf, and even the abyssal plain, all have characteristic oligochaetes associated with them. Four families of oligochaetes occur along the east coast of North America, namely, the Mega- scolecidae (with 2 species), Naididae (8 spe- cies), Enchytraeidae (13 species), and Tubifici- dae (22 species) ; members of the first named family are known as megadriles and the last three as microdriles (to which much of the fol- lowing discussion applies). Definition and Diagnostic Characters Oligochaeta are typically vermiform, cylindri- cal, segmented, bilaterally symmetrical, her- 1 National Museum of Natural Science, Ottawa, Canada, and Systematics-Ecology Program; present address: Great Lakes Biolimnology Laboratory Canada Centre for Inland Waters, 867 Lakeshore Road Burling- ton, Ontario, Canada L7R 4A6. 2 Department of Zoology, University of Toronto, Toronto 5, Ontario, Canada, and Systematics-Ecology Program; present address: Fisheries Research Board of Canada, Biological Station, St. Andrews, New Bruns- wick, Canada. maphroditic annelids with a spacious coelom, a prostomium, an anterior ventral mouth, and a posterior anus. Conventionally segments are numbered in Roman numerals, beginning at the anterior end with the peristomium as segment I. Each segment, except the persistomium, usually bears four bundles of setae, two dorsolateral, and two ventrolateral (shortened to dorsal and ventral in the key) which are implanted directly in the body wall; parapodia are absent. There are two basic types of setae whose number and morphology in the various body regions are taxonomically important: (a) crochets, which can be straight, curved or sigmoid, may or may not possess a more or less median thickening (the node or nodulus) and which may have rounded, simple-pointed or bifid (forked) distal ends; crochets are found in all oligochaete fam- ilies; (b) hair setae, which are elongate, simple- pointed structures, without a node, and whose surface may be smooth, finely serrated, or in- vested with a number of very fine lateral hair- lets; hair setae are found only in the dorsal bundles of some Naididae and Tubificidae among the families with marine species. At sexual ma- turity the epidermis of a few anterior segments is thickened, forming a dorsolateral or annular clitellum which secretes the cocoon containing one to several eggs. Oligochaetes do not have a larval stage, and juveniles resembling small adults emerge from the cocoons after a few weeks or months of development. A double ganglionated ventral nerve cord extends through the length of the body: anteriorly it divides clitellum Figure 1. — A microdrile. and passes upwards on either side of the pharynx to, or just posteriorly to, the peristomium where it joins a dorsal bilobed cerebral ganglion (or brain). The gut begins as a short narrow esophagus, then widens into a simple tube (con- stricted by the septa dividing each segment) which extends throughout the body length. Just posteriorly to the esophagus, the Enchytraeidae, Naididae, and Tubificidae (microdriles) possess a thickened pad of cells dorsally (the pharynx) which can be extruded through the mouth and to which, presumably, food particles adhere. In the Megascolecidae (megadriles) a muscular thickening of the gut wall, the gizzard, is usually situated near the anterior end of the esophagus. In the microdrile families bundles of darkly staining secretory cells, the pharyngeal glands, are found associated with the septa and gut in the region of segments III to VII. In Tubificidae and Naididae these glands are usually diffuse cell masses located laterally and dorsally, or dor- sally, to the gut. In Enchytraeidae, however, the pharyngeal glands are discrete organs with a characteristic appearance and distribution. Peptonephridia, found only in some genera of Enchytraeidae, are also glandular organs asso- ciated with the anterior part of the gut; the term is used in the sense of Nielsen and Christensen (1959) to denote paired, tubular diverticulae which arise at the esophageal-pharyngeal junc- tion. The vascular system consists of dorsal and ventral blood vessels which are connected in each segment either directly by dorsoventral commis- sures or indirectly through a blood plexus sur- rounding the gut. Excretion is by means of seg- mentary arranged paired nephridia. Each nephridium consists of a ciliated funnel which opens in the coelom, and a convoluted tubule which communicates with the exterior through a nephridiopore located ventrally on the adja- cent posterior segment. Nephridia are usually absent in a variable number of anterior seg- ments, in the region of the genitalia, in some posterior segments, and in some species they may be reduced to one, or a few, in number. The coelom is usually a simple cavity filled with coelomic fluid; in some genera and species the latter contains large numbers of coelomocytes which are large, spherical to ovoid, free cells. The genital system consists basically of male, female, and spermathecal components. The male component consists of one or two, rarely more, pairs of testes whose products (sperm) are con- ducted to the exterior by a more or less complex set of male genitalia. The sperm from each pair of testes is collected by a pair of male funnels (located on the posterior septum of the testis seg- ment) which each open into a tubular vas de- ferens. The latter usually join paired ectoder- mal storage and intromittant organs (the atria and penes), or only one of these, which open to the exterior through the male pores (see also Fig. 2 and under "microdriles" and "Megasco- lecidae" below) . The female system is composed of one or two pairs of ovaries and small female funnels which are situated ventrolateral^ just posteriad to the last testes segment. The sper- mathecal component, whose function is to store sperm after copulation, consists typically of a pair of ectodermal pouches, the spermathecae, which are located near the region of the gonads (except in the Enchytraeidae where they are // w Longitudinal / views \ SETAE — Prostomium -Peristomium ^1 Segment II - Pharyngeal gland -Hair seta Dorsal blood vessel Ventral vessel Nerve cord Gut Wv Dorsal setae Anterior sperm sac Testis *.hl^ Spermathecajp^ Clitellum — Ovary Vas deferens Atrium Penis— s Prostate Penial seta Female, funnel Brain Pharynx hair smooth curved straight lateral hairlets serrate upper tooth round simple Difid Figure 2. — Left, exterior of anterior portion of an oligochaete; middle, interior view; right, basic types of setae. teriorly to segment V). Each consists of an ampulla and a duct to the exterior through the sper- '. Sperm within the spermathecae mdom masses, or organized into ularly oriented, usually radially spermatozeugmata. In the genital e species the ventral setae may be dined as genital (penial and sper- ae. The latter can differ markedly ipe, or both, from the remaining ;omatic setae). The form of the their segmental position are of rstematic importance at all taxo- especially in the microdriles; in herefore, only sexually mature in- be identified with certainty. are all small worms (usually up liameter and 60 mm long) without usually with a variable number of bundle in different body regions. i genital setae are present in some Tubificidae, but both are absent ;idae: the dorsal chrochets of usually referred to as needle setae, les have one pair of spermathecae, ital ducts which are located in, or segment just posteriorly to the is, the vasa deferentia penetrate im (Fig. 2). The clitellum is one 1 in the smaller species it may be and confined to one segment. In :ally important tubificid male gen- . deferentia enter ectodermal atria, iair) consisting of a layer of inner nd an outer muscular layer cov- oneal epithelium. The proximal ium or atrial duct may terminate >re either simply, or, more usually, of modified penis which consists ieep infolded ring of body wall iso- al cylindrical organ; this, a true surrounded by a thickened layer vn as the penis sheath. A mass of g cells, the prostate gland, whose bably provide nutrients and bind- le sperm mass prior to copulation, ;he atrium by a stalk composed of messes of the gland cells. In some 'er, the prostate gland is a diffuse surrounding the whole, or part, of the atrium, and is absent as a discrete organ in at least one species. The Megascolecidae, which are more nearly related to the familiar lumbricid earthworms than to the previous three families, are large worms (exceeding 1.5 to 2.0 mm in diameter and 50 mm in length) , usually with a gizzard and with two setae in each bundle. The important difference between the male genitalia of mi- crodriles and megascolecids is that, whereas in microdriles the vasa deferentia pass through only one septum, in megascolecids the vasa de- ferentia always penetrate about six septa before opening to the exterior: in the case of the ma- rine species, two pairs of testes occur in seg- ments X and XI, and one pair of male openings are located on segment XVII or XVIII. Two to four pairs of spermathecal openings are pre- sent, the most posterior of which is located in or near intersegmental furrow 8/9. The clitel- lum is more than one cell thick and begins in the region of the thirteenth segment. Ecology The marine Oligochaeta are essentially benthic organisms which live within, and feed on, the bottom deposits. Many species are free-burrow- insr animals which feed indiscriminately on the bottom deposits, while others are meiobenthic (interstitial) worms which inhabit the inter- stices of the substrata and feed only on the very small organic particles, or browse off material from the surfaces of the larger particles. Mega- scolecidae and some Naididae, Tubificidae, and Enchytraeidae are free-burrowing, and some Tubificidae, Enchytraeidae, and possibly some Naididae, are meiobenthic. The major require- ments for the survival of oligochaetes in the marine environment, apart from the obvious physiological adaptions, therefore, are the avail- ability of suitable substrata and their ability to compete successfully with other deposit feeders in a given habitat (for example, Polychaeta, Echinodermata, Mollusca, Crustacea). Despite the fact that such situations are frequent from the littoral zone to the abyssal plain, the Oligo- chaeta are only now being recognized as regular components of the marine benthic ecosystem; hence little specific information on their ecology is available. It is known, however, that oligo- chaetes assume a very important role in polluted areas; habitats in which organic matter can ac- cumulate and oxygen can become depleted, are often inhabitated by large populations of a few species of oligochaetes, especially Tubificidae. For example, in the San Francisco Bay system Brinkhurst and Simmons (1968) found that the Tubificidae formed up to 97.8% of the total bot- tom fauna in grossly polluted habitats, and con- cluded that "changes in abundance of certain species may yield good supporting evidence of the nature and source of pollution materials" (p. 193). The major types of habitats in which marine or brackish water oligochaetes can be found may be summarized as follows: 1. Littoral zone. a. In damp sand and mud (Enchytraeidae, Tubificidae, Megascolecidae) . b. In and under decaying seaweed (En- chytraeidae, Megascolecidae) . c. Under stones and rocks lying on sand (Tubificidae). d. In or near sources of fresh water on the beach (Enchytraeidae, Tubificidae). 2. Sublittoral zone. a. In the sediment (Enchytraeidae, Tubi- ficidae, Naididae). b. On sediment or plant surfaces (Naidi- dae). Collecting Methods Marine Oligochaeta are collected by extracting fixed or living worms from the sediment or veg- etable material in which they live. In the lit- toral zone the substrate may be collected simply by digging or scooping material into containers, or by a simple coring device. Any of the various automatic corers, dredges or grabs, or in very shallow water a fine mesh hand net, are suitable for obtaining sublittoral material. If samples can be processed immediately to extract living worms (especially for Enchytraeidae) , the sed- iment is washed through a series of wire screens, the finest having a mesh diameter of 0.5 mm or less; all but the very small oligochaetes are re- tained by the latter. Material to be killed and fixed (before or after sorting) is treated as fol- lows: animals are narcotized in 0.015% pro- pylene phenoxetol, fixed in 10% Formalin solu- tion for 48 hr, and stored in 85% ethanol. Examination Procedure The anatomy of the oligochaetes can be studied by a variety of methods ; the keys have been de- signed so that wherever possible taxonomic characters can be observed using a minimum of manipulation or treatment. Characters used in couplets 1 to 11 can be seen by mounting fixed worms temporarily in glycerol or Amman's lactophenol (400 g carbolic acid, 400 ml lactic acid, 800 ml glycerol, 400 ml water) . The latter clears tissue by maceration and is therefore not recommended when subsequent treatment may be necessary; it is, however, a useful medium for mounting large collections for routine iden- tification (Brinkhurst, 1963). In general, ex- ternal characters and cuticular structures can be examined by simple whole mounts. From coup- let 12, however, internal characteristics assume increasing importance and are best examined using stained, whole, or dissected, animals. The following procedure has proven effective; worms are stained in acetic haematoxylin, washed, and transferred to acid alcohol (5 drops of hydro- chloric acid per 50 ml of 70% ethanol) until they are almost completely destained (to a very light red or pink color) ; the animals are then "blued" in alkaline alcohol (5 drops of concen- trated ammonium hydroxide per 50 ml of 10% ethanol) and dehydrated in 100% ethanol. One of two procedures may then follow: (a) the whole animal may be cleared in xylol and mount- ed in Canada balsam or (b) the genitalia of the worms may be dissected out using microscalpels or fine sharpened needles, and the parts cleared in xylol and mounted in Canada balsam. Most characters used in the key can be ob- served on whole stained animals, but dissection may be necessary for a critical examination of the genitalia of some species. If only one or two individuals are available for identification, it is recommended that a stained whole worm, mount- ed temporarily in xylol (xylol is highly volatile and the fumes are inflammable and toxic, there- fore great care should be exercised) should be examined briefly to decide whether any further treatment is necessary. If dissection is needed the animal is returned to 100% ethanol because xylol makes the tissues very brittle and dissection almost impossible. If living material is available, small specimens can be examined microscopically by mounting them in a drop of seawater, immobilizing them with gentle pressure on a cover slip, and blot- ting off the excess water. Larger individuals are immobilized by mounting them in a drop of gelatine solution, which is just ready to set, and placing a cover slip over the mountant using gentle pressure. Details of the alimentary canal, pharyngeal glands, and peptonephridia of the Enchytraeidae can be seen more readily in living animals. Because most marine oligochaetes are poorly known and because many have been described only within the last 6 years, the known distribu- tion of many species merely reflects the distribu- tion and collecting activities of a few specialists. Therefore, the scope of this work is expanded somewhat to include species which are known from the entire east coast of North America. It is hoped that this introduction to the marine Oligochaeta will stimulate further interest in this important class and be instrumental in amassing more information on their ecology and distribu- tion. KEY TO THE MARINE OLIGOCHAETA OF THE EAST COAST OF NORTH AMERICA The first five couplets of the key, which strictly apply only to those species included in this key, are designed so that immature Oligochaeta may be identified to their family; immature Naididae and Enchytraeidae can usually be identified to species but immature Tubificidae usually cannot. 1 Hair setae present 2 1 Hair setae absent 3 2 (1) Dorsal setae absent on segments II to V. Eyes usually present. Male and sperma- thecal pores situated on segments VI and V respectively NAIDIDAE (in part). 1 . . 9 Figure 3. — Generalized naidid; lateral view, anterior segments. 2 (1) Dorsal setae present from seg- ment II onwards. Eyes always absent. Male and spermathecal pores situated on segments XI and X respectively TUBI- FICIDAE (in part) 12 Figure 4. — Generalized tubificid; lateral view anterior segments. 3 (1) Setae with bifid ends, at least in some parts of the body. Body wall generally thin, imparting fragile appearance to worm. Genital pores on segments V and VI, or X and XI 4 3 (1) All setae with simple-pointed or rounded ends. Body wall generally thick, robust, im- parting rigid appearance to worm. Male pores on segment XII or XVIII 5 "In part" indicates that only part of a taxon will key out at that point. 4 (3) Dorsal setae absent from segments II to IV, or totally absent. Genitalia situated in segments V and VI NAIDIDAE (in part) 6 4 (3) Dorsal setae present on all segments except the peristomium. Genitalia situated in segments X and XI TUBIFICIDAE (in part) 17 5 (3) Worms exceed 1.5 to 2.0 mm in di- ameter. Setae two per bundle on all segments. Male pores situated on segment XVIII. Citellum begins on segment XIII MEGASCOLECI- DAE 45 Figure 5. — Generalized megascolecid; lateral view, anterior segments. 5 (3) Worms usually less than 1.0 mm in di- ameter. Setae very rarely two per bundle in every segment and may be totally absent in some parts of the body. Clitellum begins on segment XI ENCHYTRAEIDAE 33 Figure 6. — Generalized enchytraeid; lateral view, anterior segments. 6 (4) Dorsal setae present, beginning in segment V. Ven- tral setal bundle of segment II fan-shaped and di- rected anteriorly Paranais litoralis Figure 7. — Lateral view, anterior segments. 6 (4) Dorsal setae totally absent. Ventral setal bundle not fan-shaped 7 ^\ 7 (6) Worms commensal on gastropod molluscs, living in the mantle cavity and kidney. Setae with long strongly curved teeth (Fig. 8). .Chaetogaster limnaei Figure 8. — Somatic seta. 7 (6) Worms free-living. Setae with moderately curved distal ends (Fig. 9), Figure 9. — Somatic seta. 8 (7) Worms up to 25 mm long. Longest seta of segment II more than 200 /x long, 4.5 /x thick. Penial setae three to five per bundle Chaetogaster diapkanus 8 (7) Worms up to 7 mm long. Longest seta of segment II less than 165 /x long, 2 jx thick. Penial setae two per bundle Chaetogaster cristallinus 9 (2) Prostomium produced into long proboscis (Fig. 10) Stylaria lacustris Figure 10. — Ventral view of anterior segments, showing elongate prostomium. 9 (2) Prostomium rounded 10 10 (9) Teeth of needle setae (dorsal crochets) long and nearly parallel (Fig. 11). Upper tooth of all ventral setae about twice as long as the lower (Fig. 12) Nais elinguis Figure 11 (left). — Distal end of needle seta. Figure 12 (right). — Ventral seta. II 10 (9) Teeth of needle setae short, diverging (Fig. 13). Potserior ventral setae with approximately equal teeth (Fig. 14) 11 Figure 13 (left). — Needle seta. Figure 14 (right). — Ventral seta from a posterior segment. 11 (10) Ventral setae of segments II to V with upper tooth longer than lower; setal teeth diverge at an angle less than 45° (Fig. 15) Nais variabilis Figure 15. — Ventral seta from an anterior segment. 11 (10) Ventral setae of segments II to V similar to posterior setae, with nearly equal teeth; setal teeth diverge at an angle greater than 45° (Fig. 16) Nais communis Figure 16. — Ventral seta from an anterior segment. 12 (2) Body wall smooth. Hair setae twisted distally (Fig. 17). Vasa deferentia shorter than atria; prostate gland diffuse (Fig. 18). Coelomocytes large and very numerous. . Monopylephorus irroratus Figure 17 (top). — Lateral view of three segments from middle re- gion of body; H, hair seta. Figure 18 (bottom). — Diagrammatic lateral view of male genitalia (segment XI); P, prostate gland; V, vas deferens. 12 (2) Body wall with conspicuous papillae (Fig. 19) or with accu- mulations of foreign particles in ridges of epidermis (Fig. 20) . Hair setae not twisted distally. Vasa deferentia long- er than atria; prostate gland compact (Fig. 21). Coelo- mocytes absent 13 Figure 19 (top). — Lateral view of body wall, papillate forms. Figure 20 (middle). — Lateral view of body wall, granulate forms. Figure 21 (bottom). — Diagrammatic lateral view of male geni- talia; P, prostate gland; V, vas deferens. 13 (12) All or some posterior dorsal setae single-pointed. 13 (12) All posterior dorsal setae (crochets) bifid. . . . 14 15 14 (13) Body wall with large papillae. Dorsal crochets bifid with upper tooth reduced or rudimentary, or single-pointed, or both (Fig. 22). Ven- tral setae with upper tooth shorter than lower (Fig. 23) Peloscolex benedeni (in part) Figure 22 (left). — Dorsal crochet. Figure 23 (right). — Ventral seta. 14 (13) Body wall surrounded, at least in part, by accumulated foreign par- ticles; papillae absent. Dorsal crochets bifid with equal teeth up to about segment VI (Fig. 24); dorsal crochets from segment VII long, single-pointed, hairlike. Ventral setae with upper tooth longer than the lower (Fig. 25) Peloscolex intermedius Figure 24 (left). — Dorsal crochet, anterior segment. Figure 25 (right). — Ventral seta. 15 (IS) Body wall covered with large papillae. Posterior dorsal crochets with upper tooth shorter and thinner than the lower (Fig. 26). Anterior hair setae short and strongly bent Peloscolex dukei Figure 26. — Dorsal crochet. 15 (13) Body wall surrounded, at least in part, by accumulations of foreign particles; papillae small or absent. Posterior dorsal crochets with subequal teeth (Fig. 27, 28). Hair setae long, straight 16 Figures 27 and 28. — Dorsal crochets. 16 (15) Anterior dorsal crochets with at least one distinct intermediate tooth (Fig. 29) Peloscolex nerthoides Figure 29. — Dorsal crochet. 16 (15) Anterior dorsal crochets without intermediate teeth (Fig. 30) Peloscolex apectinatus Figure 30. — Dorsal crochet. 17 (4) Body wall covered, at least in part, by papillae or accumulation of foreign particles, or both. Penial setae absent 18 17 (4) Body wall smooth; if, rarely, some foreign particles surround body, then penial setae are present 19 18 (17) Body wall with large papillae. Dorsal crochets bifid with upper tooth reduced or rudimentary or single-pointed, or both (Fig. 31) Peloscolex benedeni (in part) Figure 31. — Range of form of dorsal crochets. 18 (17) Two or more anterior segments of body wall without papillae, or body with accumulations of foreign particles on some parts. Dorsal crochets with up- per tooth well developed; never rudimentary or absent (Fig. 32) Peloscolex gabriellae Figure 32. — Dorsal crochet. 10 19 (17) Cuticular penis sheath present. Penial setae absent 20 19 (17) Cuticular penis sheath absent. Penial setae present or absent 22 20 (19) Penis sheath long, cylindrical, with hooded end ; over eight times longer than wide (Fig. 33). . . Limnodrilus hoffmeisteri Figure 33. — Penis sheath. 20 (19) Penis sheath thimble-shaped to somewhat elongate; less than six times longer than wide. 21 Z3 21 (20) Penis sheath elongate with a cuticular hook laterally, near the distal end (Fig. 34). Anteriorly setae shal- lowly bifid (Fig. 35); posteriorly setae single-pointed (Fig. 36) Tubifex longipenis Figure 34 (top). — Penis sheath. Figure 35 (bottom left).- — Dorsal and ventral anterior setae. Figure 36 (bottom right). — Dorsal and ventral posterior setae. 21 (20) Penis sheath thimble-shaped, unornamented (Fig. 37). All setae bifid (Fig. 38) Tubifex pseudogaster Figure 37 (left). — Penis sheath. Figure 38 (right). — Dorsal and ventral setae. 22 (19) Anterior setae two per bundle, some blunt and some with trifid ends (Fig. 39). Posterior setae one per bundle, bifid with short teeth which diverge at an obtuse angle; upper tooth often reduced or rudimentary; setal node characteristically asymmetrical (Fig. 40) Clitellio arenicolus Figure 39 (left). — Anterior seta. Figure 40 (right). — Posterior seta. 22 (19) Anterior setae rarely two per bundle, never with trifid ends. Posterior setae rarely one per bundle, never with teeth diverging at an obtuse angle. Setal node more or less symmetrical 23 11 23 (22) Genital setae (penial, or spermathecal, or both) present 24 23 (22) Genital setae absent 29 24 (23) Somatic setae broad, bifid, with upper tooth shorter and much thin- ner than lower (Fig. 41). Spermathecal setae three to four per bundle, single-pointed to very faintly bifid (Fig. 42). External male pore single, median Smithsonidrilus marinus Figure 41 (left).- Figure 42 (right). ■Somatic seta. — Spermathecal seta. 24 (23) Somatic setae not of previous form. If present, spermathecal setae not of previous form. External male pores paired, ventrolateral 25 25 (24) Posterior dorsal setae one per bundle with strongly curved ends and small thin upper teeth (Fig. 43). Posterior ventral setae normal bifids, much smaller than dorsals (Fig. 44). Sperma- tozeugmata present. Spermathecal setae present Isochaeta hamata Figure 43 (left). — Dorsal seta. Figure 44 (right). — Ventral seta. 25 (24) Posterior dorsal setae more than one per bundle and not of previous form. Sperma- tozeugmata absent. Spermathecal setae absent 26 26 (25) Posterior dorsal setae single-pointed, strongly curved (Fig. 45). Penial setae of two types; each penial bundle contains one giant pe- nial and 8 to 12 small clubbed penials with small reflexed distal tooth (Fig. 46). Two prostate glands join each pear-shaped penial bulb (Fig. 47) Adelodrilus anisosetosus Figure 45 (top left). — Dorsal seta. Figure 46 (top right). — Distal end of small penial seta. Figure 47 (bottom). — Lateral view of male genitalia; A, atrium; B, penial bulb; G, giant penial setae; P, prostate gland. 26 (25) Posterior dorsal setae not as Figure 45. Penial setae of one type; penials approximately the same size as the somatic setae. Two prostate glands attached to each atrium 27 12 27 (26) Penial setae two to three per bundle, strongly curved distally (Fig. 48). Atria about five times longer than wide (Fig. 49) Phallodrilus obscurus Figure 48 (top). — Penial seta. Figure 49 (bottom). — Lateral view of male genitalia; A, atrium; P, prostate gland. 27 (26) Penial setae 7 to 13 per bundle; hooked or simple- pointed distally. Atria not of above form (? 28 28 (27) Penial setae 10 to 13 per bundle; hooked distally (Fig. 50). All somatic setae bifid. Atria about eight times longer than it is wide (Fig. 51) Phallodrilus coeloprostatus Figure 50 (top). — Penial seta. Figure 51 (bottom). — Lateral view of male genitalia; A, atrium; P, prostate. (T 28 (27) Penial setae 7 to 10 per bundle, simple-pointed distally (Fig. 52). Posterior somatic setae simple-pointed. Atria small, pear-shaped, erect (Fig. 53) Phallodrilus parviatriatus Figure 52 (top). — Penial seta. Figure 53 (bottom). — Lateral view of male genitalia; A, atrium; P, prostate; Ps, penial setae. 29 (23) Coelomocytes large and very numerous (Fig. 54). Prostate gland diffuse 30 Figure 54. — Anterior segment; C, coelomocytes. 29 (23) Coelomocytes absent, or small and few in number. Prostate gland compact, or absent. 31 13 30 (29) Length 8 to 15 mm. Spermatheca single, opening midventrally (Fig. 55). Penis absent Monopylephorus parvus Figure 55. — Dorsal view of genitalia, segments X and XI; S, spermatheca; P, prostate. 30 (29) Length 25 to 46 mm. Spermathecae paired, open- ing medially to line of ventral setae (Fig. 56). Large penis present. . . .Monopylephorus rubroniveus Figure 56. — Dorsal view of genitalia, segments X and XI; S, spermathecae; Pe, penis; P, prostate. 31 (29) Spermatheca single, opening middorsally (Fig. 57). Each atrium with two prostate glands Phallodrilus monospermathectis Figure 57. — Lateral view of genitalia, segments X and XI; S, spermatheca; P, prostate. 31 (29) Spermathecae paired; open ventrolateral. Each atrium with no, or one, prostate gland. 32 32 (31) Length about 10 mm. Setae bifid with subequal teeth (Fig. 58). Atria short, cylindrical, reflexed distally, each with a com- pact prostate gland (Fig. 59). Male and spermathecal pores in common midventral bur- sae Limnodriloides medioporus Figure 58 (top).— Seta. Figure 59 (bottom). — Lateral view of male genitalia; A, atrium; P, prostate. 32 (31) Length 30 to 65 mm. Setae bifid with upper tooth small, rudimentary or absent (Fig. 60). Atria very long, cylindrical, without an external pros- tate gland (Fig. 61). Male and spermathecal pores ventrolateral Clitellio arenarius Figure 60 (top).— Seta. Figure 61 (bottom). — Lateral view of male genitalia; A, atrium; V, vas deferens. 14 33 (5) Dorsal and ventral setae absent from all segments. Coelomocytes in posterior segments large, opaque, white flat discs (Fig. 62) Marionina achaeta Figure 62. — Lateral view of segment from middle region of body; C, coelomocyte. 33 (5) Dorsal, or ventral, or both, setae present on all, or some segments. Coelomocytes not of the above form 34 34 (33) Dorsal setae absent from all segments 35 34 (33) Dorsal setae present on all, or some, segments 38 35 (3A) Whenever present, ventral setae one per bundle 36 35 (3 %) Whenever present, ventral setae two per bundle 37 36 (35) Ventral setae present on all segments. Proximal ends of setae hook-shaped (Fig. 63). Pharyngeal glands united dorsally (Fig. 64) Lumbricillus codensis Figure 63 (top). — Ventral seta. Figure 64 (bottom). — Dorsal view of seg- ments V, VI, and 1/2 VII; A, anterior direction. (Stippled areas in this, and Fig. 66 to 70, are pharyngeal glands.) 36 (35) Ventral setae absent from about the first 15 segments. Proximal ends of setae broad, spat- ulate (Fig. 65). Pharyngeal glands not united dorsally (Fig. 66). . . .Hemigrania postclitellochaeta Figure 65 (top). — Ventral seta. Figure 66 (bottom). — Dorsal view of pharyngeal glands; A, anterior direction. y 37 (35) Ventral setae present on all segments. With three pharyngeal glands, all united dorsally (Fig. 67). ... Marionina subterranea Figure 67. — Dorsal view of pharyngeal glands; A, anterior direction. 37 (35) Ventral setae present on segments II to VI, but are absent thereafter. With two pharyngeal glands, both united dor- sally (Fig. 68) Marionina preclitellochaeta Figure 68. — Dorsal view of pharyngeal glands; A, anterior direction. 15 38 (34) One or two setae per bundle. Dorsal setae absent from segment II at least 39 38 (34) Two to nine setae per bundle (very rarely one). Dorsal setae present on all segments. 40 39 (38) Dorsal and ventral setae two per bundle; dorsal setae of segment II, and sometimes III, absent. Posterior pharyngeal glands not united dorsally (Fig. 69) Marionina southerni Figure 69. — Dorsal view of pharyngeal glands; A, anterior direction. 39 (38) Dorsal and ventral setae two per bundle on segments III to V; dorsals absent on segments II and VII to XVI; all setae one per bundle from segment XVII. Posterior pharyngeal glands united dorsally (Fig. 70) Marionina welchi Figure 70. — Dorsal view of pharyngeal glands; A, anterior direction. 40 (38) Peptonephridia (P) present (seen best in living speci- mens) (Fig. 71). Fifteen to 35 mm long. Forty-six to 65 segments 41 Figure 71. — Lateral view of anterior segments; P, peptonephridium; Ph, pharyngeal glands. 40 (38) Peptonephridia absent. Four to 15 mm long. Twenty-six to 50 segments 42 41 (40) Spermathecal duct covered by a dense layer of gland cells (Fig. 72). Twenty to 35 mm long. Two to five setae per bundle posteriorly Enchytraeus albidus Figure 72. — Lateral view of spermatheca ; S, ampulla; G, gland cells. 41 (40) Spermathecal duct devoid of glandular cells except for a ring of glands round the spermathecal opening (Fig. 73). Fifteen to 20 mm long. One or two setae per bundle posteriorly Enchytraeus capitatus Figure 73. — Lateral view of spermatheca; S, ampulla; G, gland cells. 16 42 (40) Setae distinctly sigmoid 43 42 (40) Setae straight or curved at one end 44 43 (42) Setae with a distinct node (Fig. 74). Twenty- six to 28 segments Cernosvitoviella immota Figure 74. — Seta. 43 (42) Setae without a node (Fig. 75). Thirty-eight to 42 segments Lumbricilliis lineatus Figure 75. — Seta. 44 (42) Setae straight (Fig. 76). Setal bundle fan-shaped, with setae decreasing in size towards center of bundle (Fig. 77) . Gut diverticulae present in segment VIII (best seen in living material). Thirty-nine to 50 segments Henlea ventriculosa Figure 76 (left).— Seta. Figure 77 (right). — One setal bundle. 44 (42) Setae curved distally (Fig. 78). Setal bundles not of above form. No gut diverticulae. Twenty seven to 30 segments Marionina spicula Figure 78. — Seta. ^- 45 (5) Mature worms up to 72 mm long, 3 mm diameter. Small spermathecal pores located in line with, or slightly lateral to, the lateralmost setae of the ventral bundles (Fig. 79). Dorsal setae in posterior segments in regular ranks (Fig. 80). Transverse genital marking present on ventral sur- face between segments XIX and XX. Pontodrilus bermudensis Figure 79 (top). — Ventral view of segments VII to IX; A, B, lines of ventral setae; B, lateralmost; L, seta; S, sperma- thecal pores. Figure 80 (bottom). — Dorsal view of some posterior segments; C, D, lines of dorsal setae; E, seta. VII . — L VIII \ S A B IX • • L—LL-c — - D . . ._ E 17 45 (5) Mature worms up to 180 mm long, 4 mm diameter. Small spermathecal pores located medial to (never in line with) the lateralmost setae of the ventral bundles (Fig. 81). Dorsal setae in posterior segments irregular, not in orderly ranks (Fig. 82). No transverse genital marking on segment XIX or XX Pontodrilus gracilis Figure 81 (top) . — Ventral view of segments VII to IX; A, B, lines of ventral setae; B, lateralmost; L, seta; S, spermathecal pores. Figure 82 (bottom). — Dorsal view of some posterior segments; C, D, theoretical lines of dorsal setae; E, seta. VII VIII IX . ._ L t "hk s , . . . "A • . .- . : - . . . ._ c n • •• * ;• ....■- -E . . . . ANNOTATED SYSTEMATIC LIST The following check list of Oligochaeta is ar- ranged systematically in families, with genera arranged alphabetically under their family and species under their genus. Notes on habitat, ecology, life histories, and distribution in east- ern North America are given where known. References to important papers are cited under families when applicable (monographs), and individual species (systematics, ecology, and distribution). References to species are cited at the end of the annotation on each species. Class OLIGOCHAETA Family NAIDIDAE. Asexual reproduc- tion, by means of fission, occurs commonly in this family. References: Sperber (1948), Brinkhurst and Jamieson (1971). Chaetogaster cristallinus Vejdovsky, 1883. Pen- sylvania. Found in brackish water. Five segments are formed anteriorly on fission. (Brinkhurst, 1964.) Chaetogaster diaphanus (Gruithuisen, 1828). Massachusetts to Georgia. Brackish water. Predatory on smaller worms. Five seg- ments formed anteriorly on fission. (Brink- hurst, 1964.) Chaetogaster limnaei von Baer, 1827. New England, but probably a cosmopolitan spe- cies. Brackish water. Commensal on pul- monate molluscs, living sometimes in the kidney but usually in the mantle cavity; feed on cercariae. (Brinkhurst, 1964.) Nais communis Piguet, 1906. Massachusetts to Georgia (?). Brackish water. Five seg- ments formed anteriorly on fission. ( Brink- hurst, 1964.) Nais elinguis Mtiller, 1773. Massachusetts to Pennsylvania, but probably a cosmopolitan species. Brackish water. Among floating seaweed. Swims actively. Five segments formed anteriorly on fission. (Brinkhurst, 1964; Lasserre, 1966.) Nais variabilis Piguet, 1906. Connecticut to Georgia (?). Brackish water. Swims ac- tively with spiral movements. Five seg- ments formed anteriorly on fission. (Brink- hurst, 1964.) Paranais litoralis (Mtiller, 1784). Nova Scotia to New Jersey. Brackish water and ma- rine. Intertidal and subtidal to 7 m. In sand, in sheltered localities. Four segments formed anteriorly on fission. (Brinkhurst, 1964; Lasserre, 1966.) Stylaria laciistris (Linnaeus, 1767). Pennsyl- vania and New Jersey, but probably a cos- mopolitan species. Brackish water. Prob- ably among vegetation. Swims actively. 18 Five segments formed anteriorly on fission. (Brinkhurst, 1964.) Family TUBIFICIDAE. References: Brinkhurst, 1963; Brinkhurst and Jamie- son, 1971. Adelodrilus anisosetosus Cook, 1969. Cape Cod Bay, Mass. Subtidal, 10 to 21 m depth. In coarse sands. (Cook, 1971.) Clitellio arenarius (Midler, 1776). Nova Scotia to Maryland, but probably a cosmopolitan species. Intertidal. In sand and gravel, often under stones and rocks. (Moore, 1905; Brinkhurst, 1965.) Clitellio arenicohis (Pierantoni, 1902). North Carolina and Florida. Subtidal, 5 to 130 m depth. (Brinkhurst, 1966.) Isochaeta hamata (Moore, 1905). Acushnet River, New Bedford, Mass. Brackish water. Intertidal, under stones. (Moore, 1905.) Limnodriloides medioporus Cook, 1969. Massa- chusetts. Subtidal, 7 to 97 m depth. In fine sands and silt. (Cook, 1971.) Limnodrilus hoffmeisteri Claparede, 1862. Massachusetts to Maryland, but probably cosmopolitan. A predominantly freshwater species sometimes found in brackish water. (Moore, 1905.) Moyiopylephorus irroratus (Verrill, 1873). Mas- sachusetts. Brackish water; intertidal freshwater seepages, often with decaying vegetable matter. (Moore, 1905; Brink- hurst, 1965.) Monopylephorus parvus Ditlevsen, 1904. Mas- sachusetts. Brackish water and marine. Intertidal, beneath stones, to about 7 m depth. (Moore, 1905.) Monopylephoms rubroniveus Levinsen, 1883. Massachusetts. Brackish water and marine. Intertidal, just beneath surface; often as- sociated with decaying vegetable matter or freshwater seepages. (Moore, 1905.) Peloscolex apectinatus Brinkhurst, 1965. Nova Scotia and Massachusetts. Subtidal, to 21 m depth. In sand. (Brinkhurst, 1965; Cook, 1971.) Peloscolex benedeni (Udekem, 1855). Nova Scotia to Connecticut, probably a cosmopol- itan species. Brackish water and marine. Intertidal, near high water line, to 18 m depth. In gravel and sand, often beneath stones in intertidal zone. (Moore, 1905; Brinkhurst, 1965; Cook, 1971.) Peloscolex dukei Cook, 1970. North Carolina. Subtidal, 19 to 200 m depth. In sand and shell fragments. (Cook, 1970a.) Peloscolex gabriellae Marcus, 1950. Massachu- setts, but probably amphi-American. A very broadly defined and problematical spe- cies of uncertain specific limits; as it is presently defined the species is found from the low water line to abyssal depths, and in brackish water. In sand. (Cook, 1970b.) Peloscolex intermedins Cook, 1969. Massachu- setts. Subtidal, 7 to 300 m depth. In very fine sands and silt. (Cook, 1971.) Peloscolex nerthoides Brinkhurst, 1965. Massa- chusetts. Subtidal, to 42 m depth. In very coarse sand. Also found in brackish water on the west coast. (Cook, 1971.) Phallodrilus coeloprostatus Cook, 1969. Cape Cod Bay, Mass. Subtidal, to 51 m depth. In sand. Mature individuals have been found in all seasons of the year. (Cook, 1971.) Phallodrilus monospermathecus (Knollner, 1935). Massachusetts, but possibly a cos- mopolitan species. Brackish water. Inter- tidal, in sandy beaches, especially in or near freshwater seepages (personal observa- tions.) Phallodrilus obscurus Cook, 1969. Cape Cod Bay, Mass. Subtidal, to 8.5 m depth. In sand. Mature individuals found in Novem- ber only. (Cook, 1971.) Phallodrilus parviatriatus Cook, 1971. Cape Cod Bay, Mass. Subtidal, to 18 m depth. In sand. Mature individuals found in Jan- uary and June. (Cook, 1971.) Smithsonidrilus marinus Brinkhurst, 1966. North Carolina and Florida. Subtidal, to 130 m. (Brinkhurst, 1966.) Tubifex longipenis Brinkhurst, 1965. Maine and Massachusetts. Subtidal, 6 to 51 m depth. In sand and coarse sand. In den- sities up to 2300 per square meter. Mature individuals found in August and September only; cocoons probably overwinter and hatch the following spring. (Cook, 1971.) Tubifex pseudogaster (Dahl, I960). Nova Scotia to Massachusetts. Brackish water. (Brink- hurst, 1965.) 19 Family ENCHYTRAEIDAE. Reference: Nielsen and Christensen, 1959. Cernosvitoviella immota (Knollner, 1935). Mas- sachusetts. In salt marshes, above high tide line. (Lasserre, 1971.) Enchytraeiis albidus Henle, 1837. Maine to North Carolina, but probably a cosmopoli- tan species. A very tolerant species whose marine habitat is usually decaying- seaweed, or under stones, near the high tide line. Also inhabits salt marshes, compost heaps, sewage beds and effluents. (Moore, 1905; Welch. 1917; Lasserre, 1971.) Enchytraeiis capitatus Btilow, 1957. Massa- chusetts and North Carolina. In seaweed deposits, and meiobenthic on the upper shore of sandy beaches. (Lasserre, 1971.) Hemigrania postclitellochaeta (Knollner, 1935). Massachusetts and North Carolina. Sub- tidal, 6 to about 70 m depth. In coarse sand. (Lasserre, 1971.) Henlea ventriculosa (Udekem, 1854). Massa- chusetts and North Carolina. Above high tide line, in salt marshes. (Lasserre, 1971.) Lumbricillus codensis Lasserre, 1971. Cape Cod Bay, Mass. Subtidal, 6 to 16 m depth. In sand. (Lasserre, 1971.) Lumbricillus lineatus (Muller, 1774). Maine to Massachusetts. Upper littoral zone. In salt marshes and seaweed deposits. (Moore, 1905; Welch, 1917; Lasserre, 1971.) Marionina achaeta Lasserre, 1964. Massachu- setts and North Carolina. Intertidal; me- iobenthic, on upper shore and at ground water level on lower shore. (Lasserre, 1971.) Marionina preclitellochaeta Nielsen and Chris- tensen, 1963. Massachusetts and North Carolina. Intertidal; meiobenthic in sand at ground water level on middle to lower shore. (Lasserre, 1971.) Marionina southern! (Cernosvitov, 1937). Mas- sachusetts and North Carolina. Intertidal. In seaweed deposits. (Lasserre, 1971.) Marionina spicula (Leuckart, 1847). Massa- chusetts and North Carolina. Intertidal. Meiobenthic in sand, and in seaweed depo- sits. (Lasserre, 1971.) Marionina subterranea, (Knollner, 1935). Mas- sachusetts and North Carolina. Lower in- tertidal. Meiobenthic in sand at ground water level. (Lasserre, 1971.) Marionina welchi Lasserre, 1971. Cape Cod Bay, Mass. Subtidal, to about 15 m depth. In sand. (Lasserre, 1971.) Family MEGASCOLECIDAE. Pontodrihis bermudensis Beddard, 1891. Vir- ginia. Above high tide line, and intertidal, on sandy beaches and under decaying sea- weed. The record of this species is cited in Wass (1965) and is apparently based on specimens identified by J. P. Moore prior to 1931; it is possible that these are refer- able to P. gracilis. Both species are keyed out, however, for the sake of completeness. Pontodrihis gracilis Gates, 1943. Florida and Beaufort, N.C. (latter record based on a personal communication by E. C. Haff, Uni- versity of Georgia). Intertidal, on sandy beaches; also banks of rivers where water is brackish. (Gates, 1943.) SELECTED BIBLIOGRAPHY BPJNKHURST, R. 0. 1963. Taxonomical studies on the Tubific- idae (Annelida, Oligochaeta). Int. Rev. Gesamten Hydrobiol. Syst. Beih. 2:1- 89. 1964. Studies on the North American aquatic Oligochaeta I: Naididae and Opistocystidae. Proc. Acad. Nat. Sci. Phila. 116:195-230. 1965. Studies on the North American aquatic Oligochaeta II: Tubificidae. Proc. Acad. Nat. Sci. Phila. 117:117-172. 1966. A contribution to the systematics of the marine Tubificidae (Annelida, Oligo- chaeta). Biol. Bull. 130:297-303. BRINKHURST, R. O., and B. G. M. JAMIESON. 1971. Aquatic Oligochaeta of the world. Oliver & Boyd, Edinburgh, 860 p. BRINKHURST, R. O., and M. L. SIMMONS. 1968. The aquatic Oligochaeta of San Francisco Bay system. Calif. Fish Game 54:180-194. COOK, D. G. 1969. The Tubificidae (Annelida, Oligo- chaeta) of Cape Cod Bay with a taxo- 20 nomic revision of the genera Phallodrilus Pierantoni, 1902, Lhnnodriloides Pieran- toni, 1903 and Spiridion Knollner, 1935. Biol. Bull. 136:9-27. 1970a. Pejoscolex dukei n. sp. and P. acu- leatus n. sp. (Oligochaeta, Tubificidae) from the North-West Atlantic, the latter being from abyssal depths. Trans. Am. Microsc. Soc. 88:492-497. 1970b. Bathyal and abyssal Tubificidae (Annelida, Oligochaeta) from the Gay Head - Bermuda transect, with descrip- tions of new genera and species. Deep- Sea Pves. 17:973-981. 1971. The Tubificidae (Annelida, Oligo- chaeta) of Cape Cod Bay, II: Ecology and systematics, with the description of Phallodrilus parviatriatus nov. sp. Biol. Bull. 141:203-221. DITLEVSEN, A. 1904. Studien an Oligochaeten. Z. Wiss. Zool. 77:398-480. GATES, G. E. 1943. On some American and Oriental earthworms. Pt. II. Family Megascolec- idae. Ohio J. Sci. 43:99-116. KNOLLNER, F. H. 1935. Okologische und systematische Untersuchungen fiber litorale und marine Oligochaten der Kieler Bucht. Zool. Jahrb. Abt. Syst. Oekol. Geogr. Tiere 66: 425-512. LASSERRE, P. 1966. Oligochetes marins des cotes de France. I. Bassin d'Arcachon: Systema- tique. Cah. Biol. Mar. 7:295-317. 1971. The marine Enchytraeidae (Anne- lida, Oligochaeta) of the eastern coast of North America, with notes on their geo- graphical distribution and habitat. Biol. Bull. 140:440-460. MOORE, J. P. 1905. Some marine Oligochaeta of New England. Proc. Acad. Nat. Sci. Phila. 57:373-399. NIELSEN, C. O., and B. CHRISTENSEN. 1959. The Enchytraeidae; critical revision and taxonomy of European species. Nat. Jutlandica 8-9:7-160. SPERBER, C. 1948. A taxonomical study of the Naidi- dae. Zool. Bidr. Upps. 28: 1-296. STEPHENSON, J. 1930. The Oligochaeta. Clarendon Press, Oxford, 978 p. WASS, M. L. 1961. A revised preliminary check list of the invertebrate fauna of marine and brackish waters of Virginia. Va. Inst. Mar. Sci. Spec. Sci. Rep. 24, p. 1-58. WELCH, P. W. 1917. The Enchytraeidae (Oligochaeta) of the Woods Hole region Mass. Trans. Am. Microsc. Soc. 36:119-138. 21 INDEX TO SCIENTIFIC NAMES Adelodrilus anisosetosus 12, 19 Cernosvitoviella immota 17, 20 Ch aet og aster cristallinus 8, 18 diaphanus 8, 18 limnaei 7, 18 CUtellio arenarius 14, 19 arenicolus 11, 19 Enchytraeidae 1-7, 15, 20 Enchytraeus albidus 16, 20 capitatns 16, 20 Hemigrania postclitellochaeta 15, 20 Hen lea ventriculosa 17, 20 Isochaeta harnata 12, 19 Limnodriloides medioporus 14, 19 Limnodrilus hoffmeisteri 11, 19 Lumbricillus codensis 15, 20 lineatus 17, 20 Marionina aehaeta 15, 20 preclitellochaeta 15, 20 southerni 16, 20 spicula 17, 20 subterranea 15, 20 welchi 16, 20 Megascolecidae 4, 7, 17, Monopylephorus irroratus 9, parvus 14, rubroniveus 14, Naididae 1-8, Nais communis 8, ellnguis 8, variabilis 8, Parana/is litoralis 7, Peloscolex apectinatus 10, benedeni 9, 10, dukei 10, gabriellae 10, intermedins 9, nerthoides 10, Phallodrihis coeloprostatus 13, monospei^mathecus 14, obscurus 13, parviatriatus 13, Pontodrilus bermudens-is 17, gracilis 18, Smithsonidrilus marinus 12, Stylaria lacustris Tub if ex longipenis 11, pseudogaster 11, Tubificidae 1-7, 9-14, 20 19 19 19 18 18 18 18 18 19 19 19 19 19 19 19 19 19 19 20 20 19 8, 18 19 19 19 22 ACKNOWLEDGMENTS Preparation of the "Marine Flora and Fauna of the Northeastern United States" is being coordinated by the following Board: Coordinator: Melbourne R. Carriker, Systematics- Ecology Program, Marine Biological Laboratory, Woods Hole, Mass. Advisers: Marie B. Abbott, Systematics-Ecology Program. Arthur G. Humes, Boston University Ma- rine Program, and Systematics-Ecology Program. Wesley N. Tiffney, Department of Biology, Boston University, and Systematics- Ecology Program. Ruth D. Turner, Museum of Comparative Zoology, Harvard University, and Sys- tematics-Ecology Program. Roland L. Wigley, National Marine Fish- eries Service, Biological Laboratory, Woods Hole, Mass. Robert T. Wilce, Department of Botany, University of Massachusetts, and Sys- tematics-Ecology Program. The Board established the format for the "Marine Flora and Fauna of the Northeastern United States," invites systematists to collaborate in the preparation of manuals, reviews manuscripts, and advises the Scientific Editor of the National Marine Fisheries Service. Preparation of the present manual was supported in part by Grant GB-24,832 from the National Science Foundation to the Systematics-Ecology Program. COORDINATOR'S COMMENTS Publication of the "Marine Flora and Fauna of the Northeastern United States" is most timely in view of the growing universal emphasis on environmental work and the urgent need for more precise and complete iden- tification of coastal organisms than has been available. It is mandatory, wherever possible, that organisms be identified accurately to species. Accurate scientific names unlock the great quantities of biological infor- mation stored in libraries, obviate duplication of research already done, and make possible prediction of attributes of organisms that have been inadequately studied. The present manual, "Annelida: Oligochaeta" covers the east coast of North America. Dr. David G. Cook began his systematic study of the oligochaete fauna of the east coast in September 1967, when he commenced a 2-year postdoctoral fellowship in the Systematics- Ecology Program sponsored first by the Ford Foundation and later by the National Science Foundation. During this period he prepared a number of manuscripts on this little known group. These have since been published and provided the background for the preparation of this manual. Dr. Ralph O. Brinkhurst, a specialist on the microdrile oligochaetes of the world, carried out his study of the New England aquatic oligochaetes during the summer of 1967 and 1968 as a visiting investigator in the Systematics-Ecology Program under the support of the Ford Foundation and the National Science Foun- dation. Manuals are available for purchase from the Superin- tendent of Documents, U.S. Government Printing Office, Washington, D.C. 20402. The manual so far published in the series and its cost per copy is listed below. COOK, DAVID G., and RALPH 0. BRINKHURST. Marine flora and fauna of the Northeastern United States. Annelida : Oligochaeta $0.35 23 GPO 796-611 349. Use of abstracts and summaries as communica- tion devices in technical articles. By F. Bruce Sanford. February 1971, iii + 11 PP-, 1 fig. 350. Research in fiscal year 1969 at the Bureau of Commercial Fisheries Biological Laboratory, Beaufort, N.C. By the Laboratory staff. No- vember 1970, ii + 49 pp., 21 figs., 17 tables. 351. Bureau of Commercial Fisheries Exploratory Fishing and Gear Research Base, Pascagoula, Mississippi, July 1, 1967 to June 30, 1969. By Harvey R. Bullis, Jr., and John R. Thompson. November 1970, iv + 29 pp., 29 figs., 1 table. 352. Upstream passage of anadromous fish through navigation locks and use of the stream for spawn- ing and nursery habitat, Cape Fear River N C 1962-66. By Paul R. Nichols and Darrell E.' Louder. October 1970, iv + 12 pp., 9 figs. t 4 tables. 356. Floating laboratory for study of aquatic organ- isms and their environment. By George R. Snyder, Theodore H. Blahm, and Robert J. Mc- Connell. May 1971, iii + 16 pp., 11 figs'. 361. Regional and other related aspects of shellfish consumption — some preliminary findings from the 1969 Consumer Panel Survey. By Morton M. Miller and Darrel A. Nash. June 1971, iv + 18 pp., 19 figs., 3 tables, 10 apps. UNITED STATES DEPARTMENT OF COMMERCE NATIONAL OCEANIC AND ATMOSPHERIC ADMINISTRATION NATIONAL MARINE FISHERIES SERVICE SCIENTIFIC PUBLICATIONS STAFF ROOM 450 1107 N.E. 45TH ST. SEATTLE WA 98105 PENN STATE UNIVERSITY LIBRARIES FOURTH CLASS n ADDD07E0ia5b1 OFFICIAL BUSINESS -■>. r- ' Ay- ■ it- ; ,<3KX jr