.^^^'°^Co Marine Flora and Fauna o the Northeastern United States. Tardigrada LELAND W. POLLOCK noaa NATIONAL OCEANIC AND / National Marine ATMOSPHERIC ADMINISTRATION / Fisheries Service NOAA TECHNICAL REPORTS National Marine Fisheries Service, Circulars The major responsibilities of the National Marine Fisheries Service (NMFSl are to monitor and assess the abundance and geo^phic distribution of fishery resources, lo under-^tand and predict fluctuations in the quantity 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, development and enforcement of domestic fisheries regulations, surveillance of foreign fishing off United States coastal waters, and the development and enforcement of international fishery agreemenU and policies. NMFS also assists the fishing industry through marketing service and economic analysis programs, and mortgage insurance and vessel construction subsidies. U collects, analyzes, 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. Publications that review in considerable detail and at a high technical level certain broad areas of research appear in this series. Technical papers originating in M'onomics studies and from management investigations appear m 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 lor other scientific and technical publications in the marine sciences. Individual copies may be obtained i unless otherwise noted) from D83, Technical Information Division. Environmental Science Information Center. NOAA. Washmgton. D.C. 20235. Recent Circulars are: 315. Synopsis of biological data on the chum salmon, Oncorhynchus keta iWalbaum) 1792. By Richard G. Bakkala. March 1970. iii + 89 p.. 15 figs.. 51 tables. 319. Bureau of Commercial Fisheries Great Lakes Fishery Laboratory. Ann Arbor, Michigan. By Bureau of Commercial Fisheries. March 1970. 8 p.. 7 figs. 330. EASTROPAC Atlas: Vols. 17. Catalog No. I 49.4:330/lvol.) 11 vols. Available from the Superintendent of Documents. I'.S. Government Printing Office. 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 + 23 p.. 8 figs.. 2 tables. 332. Pacific hake. (12 articles by 20 authors.) March 1970. iii + 152 p.. 72 figs., 47 tables. 333. Recommended practices for vessel sanitation and fish handling. By Edgar W. Bowman and Alfred Larsen. March 1970, iv + 27 p., 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 1%9. By the Laboratory sUff. August 1970, iii + 33 p., 29 figs.. 12 tables. 336. The northern fur seal. By Ralph C. Baker. Ford Wilke. and C. Howard Baltzo. April 1970. iii + 19 p.. 13 figs. 337. Program of Division of Economic Research. Bureau of Commercial Fisheries, fiscal year 1969. By Division of Economic Research. April 1970. iii + 29 p., 12 figs., 7 Ubles. 338. Bureau of Commercial Fisheries Biological Laboratory, Auke Bay. Alaska. By Bureau of Commercial Fisheries, June 1970. 8 p., 6 figs. 339. Salmon research at Ice Harbor Dam. By Wesley J. Ebel. April 1970. 6 p.. 4 figs. 340. Bureau of Commercial Fisheries Technological Laboratory, Gloucester. Massachusetts. By Bureau of Commercial Fisheries. June 1970, 8 p.. 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 Laboratory sUff. August 1970. iii + 24 p.. 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. ill + 22 p.. 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 p.. 28 figs., 9 Ubles. 344. Bureau of Commercial Fisheries Tropical Atlantic Biological Laboratory progress in research 1965 69. Miami. Florida. By Ann Weeks. October 1970. iv + 65 p.. 53 figs., 346. Sportsman's guide lo handling, smoking, and preserving Great Lakes coho salmon. By Shearon Dudley. J. T. Graikoski. H. L. Seagran. and Paul M. Earl. September 1970, iii + 28 p., 15 figs. 347. Synopsis of biological data on Pacific ocean perch, Sebastodes aiutus. By Richard L. Major and Herbert H. Shippen. December 1970. iii + 38 p.. 31 figs.. 11 tables. 349. Use of abstracts and summaries as communication devices in technical articles. By F. Bruce Sanford. February 1971, iii + 11 p., 1 fig. 350. Research in fiscal year 1969 at the Bureau of Commercial Fisheries Biological Laboratory. Beaufort. N.C. By the Laboratory staff. November 1970, li + 49 p.. 21 figs.. 17 Ubles. 351. Bureau of Commercial Fisheries Exploratory Fishing and Gear Research Base. F'ascagoula. Mississippi. July 1. 1967 to June 30, 1969. By Harvey R. BullLs. Jr. and John R. Thompson. November 1970, iv + 29 p.. 29 figs.. 1- table. 352. Upstream passage of anadromous fish through navigation locks and use of the stream for spawning and nursery habitat. Cape Fear River. N.C. 1962 66. By Paul R. Nichols and Darrell E. Louder. October 1970. iv + 12 p.. 9 figs.. 4 tables. 356. Floating laboratory for study of aquatic organisms and their environ- ment. By George R. Snyder. Theodore H. Blahm. and Robert J. McConnell. May 1971. iii + 16 p., 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 p., 19 figs., 3 tables, 10 apps. 362. Research vessels of the National Marine Fisheries Service. By Robert S. Wolf. August 1971. iii + 46 p., 25 figs.. 3 tables. For sale by the Superintendent of Documents. U.S. Government Printing Office, Washington. D.C. 20402. 364. History and development of surf clam harvesting gear. By Phillip S. Parker. October 1971. iv -f 15 p.. 16 figs. For sale by the Superintendent of Documents, U.S. Government Printing Office. Washington, D.C. 20402. 365. Processing EASTROPAC STD daU and the construction of vertical temperature and salinity sections by computer. By Forrest R. Miller and Kenneth A. Bliss. February 1972, iv + 17 p., 8 figs., 3 appendix figs. For sale by the Superintendent of Documents, U.S. Government Printing Office, Washington. D.C. 20402. 366. Key to field identification of andromous juvenile salmonids in the Pacific Northwest. By Robert J. McConnell and George R. Snyder. January 1972, iv + 6 p.. 4 figs. For sale by the Superintendent of Documents, U.S. Government Printing Office. Washington. D.C. 20402. 367. Engineering economic model for fish protein concentration processes. By K. K. Almenas, L. C. Durilla. R. C. Ernst. J. W. Gentry. M. B. Hale, and J. M. Marchello. October 1972, Iii + 175 p., 6 figs., 6 tables. For sale by the Superintendent of Documents, U.S. Government Printing Office. Washington, DC. 20402. 368. Cooperative Gulf of Mexico estuarine inventory and study. Florida: Phase I. area description. By J. Kneeland McNulty. William N. Lindall. Jr.. and James E. Sykes. November 1972, vii + 126 p., 46 figs., 62 tables. For sale by the Superintendent of Documents, U.S. Government Printing Office. Washington, D.C. 20402. 369. Field guide to the anglefishes (Pomacanthidael In the western Atlantic. By Henry A. Feddern. November 1972. iii + 10 p . 17 figs.. For sale by the Superintendent of Documents, U.S. Government Printing Office. Washington. D.C. 20402. Continued on inside back cover. NOAA Technical Report NMFS CIRC- 394 Marine Flora and Fauna of the Northeastern United States. Tardigrada LELAND W. POLLOCK SEAHLE. WA MAY 1976 UNITED STATES / NATIONAL OCEANIC AND / National Marine DEPARTMENT OF COMMERCE / ATMOSPHERIC ADMINISTRATION / Fisheries Service Elliot L. Rictiardson. Secretary/ Robert M White Administrator / Robert W Schoning. Director For Sale by the Superintcndcnl of Documents, U.S. Governmcm Printing Office Washington. D.C. 20402 Steele No. 033-017-00369-8 FOREWORD This issue of the "Circulars" is part of a subseries entitled "Marine Flora and Fauna of the North eastern 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 laxa 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 manual will be based primarily on recent and ongoing revisionary systematic re.search and a fresh examination of the plants and animals. Each major taxon. treated in a separate manual, will include an introduction, illustrated glo.ssary, 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 systematic index. These manuals are intended for use by biology students, biologists, biological oceanographcrs. 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. Whenever 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 published, the manuals will be revised, grouped, and issued as special volumes. These volumes 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 P>ology 2 Colloctinf^ methods 2 Key to the species of marine Tardigrada of the world 3 Annotated systematic list of marine Tardigrada of the world 21 List of marine Tardigrada reported from the northeastern United States 22 Selected bibliography 22 Index to scientific names 24 Acknowledgments 25 Coordinator's comment 25 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. Marine Flora and Fauna of the Northeastern United States. Tardigrada LELANDW. POLLOCK' ABSTRACT The manual includes an introduction to the general biology, an Ulustrated key, an annotated systematic list, a selected bibliography, and an index to the Tardigrada of the marine coastal areas of the world to a depth of 5,000 m. INTRODUCTION The Tardigrada (tardus. L. slow; gradus, L. step) comprise a phylum of microscopic metazoa (usually less than 1 mm in length) of uncertain phylogenetic placement. Con- siderations of their status have been based on 1) their growth by molting; 2) absence of ciliated epithelium: 3) presence of a spacious pseudocoelom in adults; 4) muscu- lature in bandlike bundles; 5) metameric, or at least repetitive, arrangement of unjointed legs, as well as of portions of the ventral nervous system and muscular system; 6) the presence of coelomocytes; 7) the absence of circular muscles; 8) a tripartite foregut; 9) a nonchitinous cuticle; and 10) the occurrence of eutely or cell constancy (although this recently has been disputed, Bertolani 1970). Most of these characteristics suggest an organizational complexity approaching that of the aschelminth phyla, especially the Rotifera and Nematoda. Characters suggest- ing relationship with the Arthropoda include the first six characters listed above; in addition, their "ladder-type" ventral nervous sytem recalls the annelid-arthropod line. Tardigradan embryology however apparently includes a total but irregular cleavage pattern and enterocoelous formation of a series of eoelomic pouches, of which only the gonocoel is retained in the adult. While this pattern of development is unlike any other known group, it is most similar to that of the deuterostomous invertebrates. Lack of clarity regarding their phylogenetic affinities suggests that the Tardigrada are far removed from their nearest phyletic neighbor. Apparently they are an old group which has become highly specialized for life in peculiar habitats, such as the water films surrounding lower plants and lining interstitial spaces between grains of sand. Morphological diversity among marine tardigrades attests to their age. On the other hand, the comparative uniformity in appearance and simplicity in morphological characters of freshwater forms supports the hypothesis that marine tardigrades are primitive. There are 43 described species of marine tardigrades included in 17 genera. Most are members of the interstitial meiofauna of sandy sediments. Since one-half of these are in mono- or ditypic genera, and two thirds have been discovered since 1950, it is likely that many more species will be described in the future. Marine tardigrades rarely exceed 0.5 mm in length and are all similar in general body plan (Fig. 1). They possess as many as 11 cephalic appendages, including lateral cirri (a), clavae (cl), external cephalic cirri (ec), internal cephalic cirri (ic), and a median cephalic cirrus (mc). Their bodies usually are cylindrical, with four pairs of legs which terminate in claws, toes, or both. These terminal appendages, the spines or papillae on the legs, and the conformation of the caudal appendage (if present) are important taxonomically. Likewise, the presence and location of somatic cirri, especially posterior- lateral cirrus (e) (Fig. 1), can be of taxonomic significance. In the order Eutardigrada, the cephalic appendages toe & claw somatic spine cirrus e 'Department of Zoology, Drew University, Madison, NJ 07940. gonopore anus caudal appendage Figure 1 . — Diagrammatic illustration of a composite marine tardigrade. Cephalic appendages: a - lateral cephalic cirrus: cl - clava: ec - external cephalic cirrus; ic - internal cephalic cirrus: mc ■ median cephalic cirrus. number and relative sizes of placoids or rod-shaped concretions within the bulbous muscular pharynx can be diagnostic. Members of at least six genera {Archechiniscus. Batillipes. Coronarctus, Echiniscoides, Parastygarctus, and Stygarctus) display sexual dimorphism in the shape and location of gonopores. Females possess rosette gonopores located midventrally at considerable distance anterior to the anus. In males, the gonopore is circular or tubular, mid-ventral and only slightly anterior to the anus. In members of the genus Halechiniscus, dimorphism is exhibited in the relation of the length of clavae relative to the lateral cirri. In males, clavae are longer than lateral cirri, while the opposite is true of females. Sex determi- nation in other marine tardigrades is based on presence of mature gametes in the gonad or on the fact that males possess two vasa deferentia while females have a single oviduct. Tardig^ada develop directly. Excepting their dispropor- tionately longer cephalic appendages and their reduced number of claws per leg, juveniles resemble miniature adults. Growth in Tardigrada is accomplished through periodic molting of all cuticular structures, including the linings of the foregut and hindgut. Apparently internal fluid pressure is reduced enough to permit defecation, oviposi- tion, and sperm penetration only during an intermolt period. Other aspects of the morphology and anatomy of marine tardigrades lie beyond the scope of this presentation. Interested readers are referred to monographs by Marcus (1936), Rudescu (1964), and Ramazzotti (1972), and to a recent review by Renaud-Mornant and Pollock (1971). ECOLOGY In recent years, ecological studies of marine Tardigrada have focused largely on those living interstitially among grains of sand (Renaud-Debyser 1959a; Schmidt 1968, 1969; Pollock 1970c; Lindgren 1971). Tardigrades are found throughout portions of intertidal beaches which undergo periodic drainage and replenishment of interstitial water. Most interstitial meiofauna, including tardigrades, are absent or uncommon in beaches of fine sand (mean grain diameter less than 300 /im) and in beaches of larger grain- size but where fine silt and debris clog pore spaces and restrict circulation. Tardigrada occupy specific portions of littoral beaches creating patterns of zonation both horizontally along the beach surface and vertically within the sediment. A "typical pattern" of species composition and distribution on a single beach becomes evident from studies of temperate, quartz sand beaches. An abundant species and from one to several less common species of Batillipes occupy superficial sand (occurring somewhat deeper in beaches under the influence of heavy surf). The abundant Batillipes dominates mid-beach sand while other Batillipes often are relegated to more landward or seaward locations. A comparatively denser concentration of Stygarctus often occurs deeper within the beach approaching the deepest sediments undergoing tidal drainage of interstitial water. Less frequently, marine tardigrades are reported from other habitats. Sublittoral specimens have been collected to a depth of 4,700 m (Renaud-Mornant 1974). Their comparative scarcity in most deepwater surveys suggests either that Tardigrada are less successful here than are many other meiofaunal groups or that sampling and/or observational techniques commonly used fail to include such small members of the meiofauna. Certain Tardigrada occur on seaweed ranging from intertidal Enteromorpha and Lichinia to offshore Sargassum. Among the several Tardigrada reportedly living ectocommensally with various other marine invertebrates, Tetrakentron synaptae Cuenot, 1892 alone possesses obvious morphological adaptations to such a life style and has been found exclusively in such a relationship. COLLECTING METHODS A complete discussion of techniques for working with interstitial meiofauna generally and marine Tardigrada specifically may be found in Hulings and Gray (1971). Quantitative extraction of tardigrades from sand requires rigorous procedures since most species are strongly thigmotactic and vigorously resist dislodgment. Anestheti- zation by flooding a small sand sample with 3.5% MgCl2 may be effective for removing Tardigrada from sediments gathered in areas of low to moderate wave activity; however, this technique is not effective quantitatively on samples from "high-energy" beaches (Gray and Rieger 1971). Soaking small quantities of sand (e.g., 10 cm' or less) in 10 times that volume of 3.5% ethanol is more effective for anesthetization. This can be followed by three or more rinses of seawater to provide revived and apparently unharmed Tardigrada quantitatively. Marine Tardigrada can be preserved well in either 5-7% neutralized Formalin or in 70% ethanol. McGinty and Higgins (1968) described a widely used technique for mount ing marine tardigrades. Specimens preserved in 7% Formalin are transferred to a 1:10 glycerin-Formalin solution which then is allowed to evaporate to glycerin (a glycerin-alcohol solution works for specimens preserved in alcohol). Tardigrades prepared by this technique can be mounted in glycerine, glycerine jelly, or Hoyer's medium. Phase contrast microscopy is necessary for fine observa- tions, especially if Hoyer's medium is used. KEY TO THE SPECIES OF MARINE TARDIGRADA OF THE WORLD The following key is designed for the artificial separation of marine tardigrades. Morphological characters are utilized for easy identification and are not intended to fully describe the animals. While examination of living animals at high power or oil immersion is imperative for complete and accurate descriptions, specimens fixed in 10% Formalin or 70% alcohol are usually recognizable. The illustrations are variously modified from original illustrations or descriptions. Important distinguishing features are indicated on each figure by short pointer lines. 1 Legs terminate in claws which attach directly, or if on toes are longer than toes 2 1 Legs terminate in toes without claws or with claws shorter than toes 12 2 1/1 Central two claws on each leg bear hairlike extensions 3 2 {!) Claws without hairlike extensions 6 3 (2) Caudal spikes absent; anterior margin of head deeply sculptured , 4 3 (2) Caudal spike present; anterior margin of head much less deeply sculptured 5 4 iS) Lateral extensions of dorsal plates end in two points Parastygarctus higgiiisi 4 {S) Lateral extensions of dorsal plates end in single point Parastygarctus sterreri 5 (5) Somatic spines on mid posterior border of somatic plate II present; 2 cusps only along margin of cephalic plate Stygarctus bradypus 5 (3) Somatic spines on somatic plate II absent; 4 cusps along margin of cephalic plate Stygarctus granulatus 6 (2) Cephalic appendages absent 7 6 (2) Cephalic appendages present 9 7 {6) First pharyngeal macroplacoid longer than second; inner diam- eter of pharyngeal tube 3 (im 8 7 {6) First macroplacoid shorter than second; inner diameter of pharyngeal tube 1.5 fim Hypsibius stenostomus 8 ( 7) Second macroplacoid shorter than third Hypsibius appelloefi 8 (7) Second macroplacoid as long as or longer than third Hypsibius geddesi 9 [6) More than four claws (usually 5-11) on each leg; distinct dorsal cutieular plates absent Echiniscoides sigismundi 9 {6) Four claws or less per leg 10 10 (5) Four claws per leg present on anterior three pairs but only three claws per leg on posterior pair; distinct dor- sal plates absent; median cirrus absent . . .Anisonyches diakidius 10 {9) Four claws per leg present on all legs 11 11 (10) Median cirrus present; distinct dorsal cuticular plates absent Coronarctus tenellus 11 {10) Median cirrus absent; dorsal cuticular plates present Echiniscus (E. quadrispinosus is the only species reported from marine environment.) 12 (i) Four to six toes without claws on each leg 13 12 (/) Four toes with claws on each leg 27 13 {12) Toes end in disc expansions 14 13 {12) Toes end in narrow lobate expansions 26 14 {IS) Projecting caudal appendage absent 15 14 {13) Projecting caudal appendage present 16 15 [H) Caudal end swollen cephalic append- ages long BatiUipes tubematis 15 (i-4) Caudal end relatively flat; cephalic appendages short 16 iH) Caudal appendage basically a single spike 16 iU) Caudal appendage more than one spike . . . 17 23 17 (iff) Caudal spike terminates in a membranous bag BatiUipes buUacaiidatus 17 [16) Caudal spike blunt or sharp tipped 18 18 (i 7) Distinct constrlction(s) present on clavae 19 18 [17) Clava uniform in width, constrictions absent 21 19 [18) Single constriction present on clava 20 19 ( 18) More than one constriction present on clavae BatiUipes annulatus 20 {19) Caudal spike from twolobed base BatiUipes gilmartini 20 (19) Caudal spike from single-Iobed base BatiUipes pennaki 21 US) Leg spines on hindmost legs short; leg spines present on anterior three pairs of legs BatiUipes minis 21 (18) Leg spines on hindmost legs long; spines absent on anterior three pairs of legs . 22 22 (21) Caudal appendage a long, slender, sharp-tipped spike . Batillipes camonensis 22 {21) Caudal appendage, a short, thick, blunt-tipped spike Batillipes similis 23 {16) Caudal appendage two-spiked Batillipes dicrocercus 23 {16) Caudal appendage bears three or more spikes 24 10 24 (23) Caudal appendage bears three spikes only 25 24 {23) Cauda! appendage bears more than three spikes; central spike longest Batillipes pkreaticus 25(24) Spikes of equallength 25 {2Jf) Lateral spikes shorter than central spike Batillipes friaufi 11 26 [13) Papilla present on fourth pair of legs; clava slightly constricted Orzeliscus belopus 26 {13} Papilla absent on fourth pair of legs; clava uniform in width Orzeliscus septentrionalis 27 (12) Middle two toes on each foot much longer than outer two toes Archechinvscus marci 27 {12) Toes approximately equal in length 28 12 28 (27) Each claw has more than two exposed points 28 {27) Each claw has one or two exposed points ... 29 30 29 (28) Body flattened dorsovenlrally; cephalic appendages reduced; ectocommensal Tetrakentron synaptae 29 {28) Body cylindrical 30 30 (29) Each claw has three exposed points 31 30 (25) Each claw has four exposed points Bathyechiniscus tetronyx 13 31 (50) Cirri e and clavae both present 32 31 [SO) Cirri e and clavae both absent Styraconyx haphceros 32 {31) Clavae long (30(im); eyes present Styraconyx paulae 14 32 iSl) Clavae short (13(im); eyes absent Styraconyx sargassi 33 (28) Body covered dorsally by tubules and gelati- nous coating Actiimrctus doryphorus 33 (2S) Body smooth, lacking tubules and gelatinous coating 34 34 (SS) Cuticle forms two caudal projections (branched or unbranched) extending more than body length 34 (33) Caudal projections less than body length or absent .35 36 15 35 [SJt) Caudal projections branched Tanarctus arborspinosus 35 iSi) Caudal projections unbranched Tanarctus tauricus 16 36 iS4) Clava bulb-shaped; less than 0.5 length of lateral cirrus; eyes present Pleocota limnoriae 36 \3^] Clava thin, more than 0.5 leng[th of lateral cirrus; eyes absent 37 37 (36) All claws crescent shaped without "anchor spikes" 37 (36) Crescent shaped claws on middle two toes of each foot with accessory "anchor spike" embedded in toe 38 (37) Five peripheral alae or membranous sheets present 38 (37) Three peripheral alae present; caudal ala smooth, undivided; lateral alae undivided Fhrarctus salvati 11 39 [38) Caudal ala slightly sculptured, not divided; lateral alae divided Florarctus antillensis 39 [38) Caudal ala deeply sculptured and nearly divided; lateral alae divided Florarctus heimi 18 40 {37) Peripheral alae present Halechiniscus intermedius 40 [S7) Peripheral alae absent 41 41 (^0) Caudal spike prominent Halechiniscus rerruinei 41 HO) Caudal spike absent . 42 19 42 {41) Cirri e present in addition to three pairs of somatic spines Halechiniscus guiteli 42 (Al) Cirri e present but additional somatic spines absent 43 43 (-42) Enlarged papilla on shank of hindmost legs; body outline smooth Halechiniscus perfectus 43 (-42) Simple leg spine present on shank of hindmost legs; body with slight sequential ridges . . Halechiniscus subterraneus 20 ANNOTATED SYSTEMATIC LIST OF MARINE TARDIGRADA OF THE WORLD The arrangement of the following list is based on the classification suggested by Ramazzotti (1972). Original descriptions of all species are available through references in the bibliography. Where subsequent descriptions are useful, they are cited parenthetically. Finally, the most typical habitat and geographical range is indicated for each species. Order HETEROTARDIGRADA Suborder Arthrotardigrada FAMILY HALECHINISCIDAE Halechiniscus guiteli Richters 1908. Interstitial, sandy intertidal. English Channel, eastern Atlantic, Mediter- ranean, Black Sea. Halechiniscus intermedins Renaud-Mornant 1967b. Inter- stitial, subtidal, coralline sand at 4 m depth. Southern Pacific. Halechiniscus perfectus Schulz 1955. Interstitial, sandy intertidal and subtidal at 170 m depth. North Sea, Medi- terranean, southern Pacific, Indian Ocean. Halechiniscus remanei Schulz 1955 (McGinty 1969). Inter- stitial, sandy intertidal to subtidal at 150 m depth. Medi- terranean, eastern Atlantic, eastern Pacific. Halechiniscus subterraneus RenaudDebyser 1959b. Inter- stitial, intertidal in coralline sand. Western Atlantic. Pleocola limnoriae Cantacuzene 1951. Commensal of isopod Limnoria lignorum and sandy intertidal to subtidal at 130 m depth. English Channel, western Atlantic. Actinarctus doryphorus Schulz 1935 (Grell 1937). Intersti- tial, sandy intertidal to subtidal at 170 m depth. One report of possibly ectocommensal relationship with Echinocyamus pusiUus (Grell 1937). North Sea, English Channel, eastern Atlantic. Tetrakentron synaptae Cuenot 1892. Ectocommensal on tentacles of holothurian, Leptosynapta galliennei. Eng- lish Channel. Styraconyx haploceros Thulin 1942. Lichens near high tidal line. English Channel. Styraconyx paulae Robotti 1971. Shallow subtidal, epizoic on coral. Styraconyx sargassi Thulin 1942. Pelagic on algae, especially Sargassum. Western Atlantic, Mediterra- nean, Gulf of Mexico, eastern Pacific. Bathyechiniscus tetronyx Steiner 1926. Questionable de- scription based on single observation from mud at 400 m depth in south Atlantic. Flararctus antiUensis Van der Land 1968 (Renaud-Mornant 1971). Interstitial in coralline sand, 3 m depth. Carib- bean. Florarctus heimi Delamare Deboutteville and Renaud- Mornant 1965 (Delamare Deboutteville and Renaud- Mornant 1966: Renaud-Mornant 1967b). Interstitial in coralline sand. Southern Pacific. Florarctus salvati Delamare Deboutteville and Renaud- Mornant 1965 (Delamare Deboutteville and Renaud Mornant 1966: Renaud Mornant 1967b). Interstitial in coralline sand. Southern Pacific. Tanarctus arborspinosus Lindgren 1971. Interstitial, inter- tidal. Western Atlantic. Tanarctus tauricus Renaud-Debyser 1959b. Interstitial, intertidal. Western Atlantic. FAMILY BATILLIPEDIDAE Batillipes acaudatus Pollock 1971. Interstitial, sandy inter- tidal. Northestern Atlantic. Batillipes annulatus DeZio 1962. Interstitial, sandy inter- tidal. Mediterranean. Batillipes huUacaudatus McGinty and Higgins 1968 (Pollock 1970a). Interstitial, sandy intertiday. Northwestern and western Atlantic. Batillipes camonensis Fize 1957. Interstitial, sandy inter- tidal. Mediterranean, Indian Ocean. Batillipes dicrocercus Pollock 1970a. Interstitial, sandy intertidal. Northwestern Atlantic. Batillipes friaufi Kiggin 1962. Interstitial, sandy intertidal. Western Atlantic. Batillipes gilmartini McGinty 1969. Interstitial, sandy intertidal. Eastern Pacific. Batillipes littoralis Renaud-Debyser 1959a. Sandy inter- tidal, interstitial. Eastern Atlantic. Batillipes minis Richters 1909 (Marcus 1927; Rudescu 1964: McGinty and Higgins 1968: Pollock 1970a, b). Sandy in tertidal and shallow subtidal, interstitial. Baltic, North Sea, Black Sea, Mediterranean, Atlantic, Indian Ocean. BatiUipes pennaki Marcus 1946 (DeZio 1962: Pollock 1970a, b). Interstitial, sandy intertidal. Northern and south- ern Atlantic, Mediterranean, Indian Ocean. 5atiMipespArea' Ellen Engett. July 1973. iv + 12 p.. 1 fig. For sale by the Superintendent of Documents, U.S. Government Printing Office. Washing ton. D.C. 20402. ;J84, Marine flora and fauna of the northeastern United States. Higher plants of the marine fringe. By Edwin T. Moul. September 1973. iii + 60 p.. 109 figs. For sale by the Superintendent of Documents. U.S. Government Printing Office. Washington. D.C. 20402, 385. Fishery publications, calendar year 1972: Lists and indexes. By Lee C. Thorson and Mary Ellen Engett. November 1973. iv + 23 p.. 1 fig. For sale bv the Superintendent of Documents. U.S. Government Printing Office, Washington. D.C. 20402. 376. Bottom-water temperatures on the continental shelf. Nova Scotia to New Jersey. By John B. Colton. Jr. and Ruth R, Stoddard. June 1973. iii + 55 p., 15 figs.. 12 appendix tables. For sale by the Superintendent of Documents. U.S. Government Printing Office. Washington, D.C. 20402. 386, Marine Flora and fauna of the northeastern United States. Pycnogo- nida. By Lawrence R. McCloskey. September 1973. iii + 12 p.. 1 fig. For sale bv the Superintendent of Documents. U.S. Government Printing Office. Washington. D.C. 20402. 377, Fishery publications, calendar year 1970: Lists and indexes. By Mary Ellen Engett and Lee C. Thorson. December 1972. iv + 34 p., 1 fig. For sale by the Superintendent of Documents. U.S. Government Printing Office. Washmgton. D.C. 20402. 387. Marine flora and fauna of the northeastern United States. Crustacea: Stomatopoda. By Raymond B. Manning. February 1974. iii + 6 p., 10 figs. For sale bv the Superintendent of Documents. U.S, Government Printing Office. Wa.shington. D.C. 20402. 378. Marine flora and fauna of the northeastern United States. Protozoa: Ciliophora. By Arthur C. Borror. September 1973. iii + 62 p.. 5 figs. For sale by the Superintendent of Documents. U.S. Government Printing Office. Washington. D.C. 20402. 388. Proceedings of the first U.S. Japan meeting on aquaculture at Tokyo. Japan. October 18 19. 1971. William N. Shaw (editor). |18 papers, 14 authors.) February 1974, iii + 133 p. For sale by the Superintendent of Documents, U.S. Government Printing Office. Washington. D.C. 20402. 379. Fishery publications, calendar year 1969: Lists and indexes. By Lee C. Thorson and Mary Ellen Engett. April 1973. iv + 31 p., 1 fig. For sale by the Superintendent of Documents. U.S. Government Printing Office. Washing- ton, D.C. 20402. 389. Marine flora and fauna of the northeastern United States. Crustacea: Decapoda. By Austin B. Williams. April 1974. iii + 50 p.. Ill figs. For sale by the Superintendent of Documents. U.S. Government Printing Office, Washington. D.C, 20402. 380, Fishery publications, calendar year 1968: Lists and indexes. By Mary- Ellen Engett and Lee C. Thorson. May 1973. iv + 24 p.. 1 fig. For sale by the Superintendent of Documents, U.S. Government Printing Office. Washing ton. D.C. 20402. .'190. Fishery publication.s. calendar year 1973: Lists and indexes. By Mary Ellen Engett'and Lee C. Thorson. September 1974, iv + 14 p.. 1 fig. For sale by the Superintendent ot Documents. U.S. Government Printing Office, Washington, D.C. 20402. MBl WHOI Librsi 5 WHSE 00517 UNITED STATES DEPARTMENT OF COMMERCE NATIONAL OCtANlC AND AIMOSPMEHIC ADMINISIHATION NATIONAL MARINE FISHERIES SERVICE SCIENTIfIC PumCATIONS STAFF ROOM 450 1107NE 45IHST SEAITIE.WA 98105 OFFICIAL BUSINESS ROStACE AND FEES PAID US DEPARTMENT OF COMMERCE COM 210 THIRD CLASS BULK RATE Marine BiolOi'jical laboratory S Library - Poriodicals Woods Hole, Ma 025^3 '2- — — /^ AMERICAS FIRST INDUSTRY