Marine Flora and Fauna of the Northeastern United States. Tardigrada
NOAA Technical Report NMFS CIRC- 394
Marine Flora and Fauna of the Northeastern United States. Tardigrada
LELAND w. POLLOCK
SE.~TTLE. IVA MAY 1976
,ct-~) t>lMOS"",(~'r cr~"~1~'\ UNITED STATES / ~ATlONAl OC[ANIC ANO N,tlOn.1 Martne ~~I f" OEPARTM~NT OF COMMERCE ~TMOSPHERIC ADMINISTRATiON / F,shertes Se'Vice ~ . ~ Elliot l. Ri~hardson. Secretary Robert M White. Admlnlst"tor / Robe'l W. Schonlng. D"eclO' 1 52 CJ' tf-":- q,."'1 _(f'....;r<,; ~'f."f"'l 0\ L
Fat S31c br th,. Sopcrintcndl'nt of Documents, U.S. GO\'l'rnmcnt Printing OffICI! W,l:5hin~1011, D.C. 20402 St(ld~ No, OB-0I7-U0369·S FOREWORD
Thisi~SUl"ofthl! "Circulars" is purl of a suhscries entitled . Marine Flora and FOllna of the North eastern United Statl!s." This subscrles will consist of original, illustrated. modern manu,'.ds on the idl'ntificat ion. dassification. and general biology of the estuarine and COilstal rTlarine planb lind animalsof lh.' northeastern lInitecStates. Manuals will bepublisbed at irregular int~rvals '''n as many taxa or lht~ rl~g-ion :~~ there arc specialists willing to col1aborate in their preparation. Tbe manuals are an outgrowth of the widely used '"Keys to Marine Invertebrates of the Woods lIole Region..' ediled hy R. I. Smith. published in 1964. and produced under the au';pices of the Systemalks·Ecology Program. Marin.., Biological Laboratory. Woods Hole. Mass. Instead of revising the '"Woods Hole Keys." the staff of the Systematics·Ecology Program decided 10 expand the gpo!-.rraphic coverage ,lOd hathymetric range and produce the kl'YS in an entire'.y new set of expandeD ~ublication.,. Tbe '·Ma.. 'ne Flora aoo F~una of thl' Northeastern United Slates" is being prepared in collaboration WIth systematic specialists in the United States and abroad. Each manual will bl' based JJrimarily on recent and onJ:oing revisionary systematic research and a fresh examinati.on of the plants and animals. Each major l:lxon. treated in a separate manual, will include an intr,)duction. illustrated glossary. uniform oriJ:inally illustrated keys, annotated check list with inform'!!ion when availahll' on distribution. bhitat. life history, and related biology. references to the majer literatuTl' of the group. and a syslcmatitO i.,dcx. These manual> are intended for use by biology studenls. biologists. biological oceanographers. informed laymen. and others wishing to identify coastal org,misms for this region. In many instances the manu,ils will serve as a guide La additional information about the species or the !iroup. Geo~.,.aphie eOVl'rage 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 OFl the continental shelf from Maine to Virginia. hut m"y vary somewhat with cae!. m"jor taxon "nd the interests of collaborators. Whenever possible representative specimens dealt with in the manu"ls y;n be deposited in reference collections of the Gray Museum, Marine Riological Laboratory. und nthcr universities and research laboratories in the region. After a -,ufficient number of manuals of related taxonomic groups have been published, the manuals wih be revised. grouped. and issuc·d as special volumes. These volumes will thus consh;t of compildiol1s of individual manu:"s within phyl" such as the Coelenterata, Arthropoda. and Mnllusca. 01' of group~ of ptlyla.
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315. Synopsis of bioloRicll1 da:a on th~ rhum saimon, Oncorhynt"hu.s kcta 3-19. Use of abstracts and summaries as communication devices in technical iWnlbauml 1792. By Rirharc G. Bi*kknln. M:uch 1970. iii + 89 p.. 15 figs •• 51 nrtirlc5. By F. Bruce Sanford. February J.971. iii + 11 p., I fig. tahles. 319. Bureau or Commercial Fisheri(>1, Great Lakes Fishery Laboru!.ory. Ann 3S0. R[!s(!:;:-<:h in (iiOcal y(!ar 19fi9 at the Bureau of Commmdal FisheriE's Arbor. Mich;y.lln. B;.· Bureau or Comlm:rcinI Fisheries. March 1970. 8 p •• 7 fiRS. Bioloh';~al Laboratory. Beaufort. N.C. By the l.aboralory staH. NGvember 1970, ii + 49 p.• 21 figs" 17 tables. 3.10. EASTROPAC Atlas: I'ols. 1·7. C.talog No. I 4~.4::I3O/{vol.J 11 vols. AV3i1able rrom the .sup£'rintl'ndent or !)ocumenl!'i. u.s. Government Printing 351. Bureau of Commercial Fisheriel> Exploratory Fishing and Gear Research Office. W:lshinl.,'1.on. D.C. 20<102. llJse, P:tsca~oula. Mississippi, July 1. 1%7 to June 30. 1969. By Harvey R. Bullis. Jr. and John R. Thompson. November IWO. iv + 2~ p.• 29 ftgn•• 1 a31. Guidelinl':' r!'ir the process-in/; or hot· smoked chub. B)' H. 1.., SeaRl"sn. J. table. T. GrJikoski. and J. A. Emer:ion. hnuar) 1970. i\' + 23 p.. 8 fiKS .. 2 table~, 352. Upstream passage or anadromous fi~h through navigation loekfi and use :J:t!. l':ttirie hakl'. 02 :trticles b)' 20 authors.) Mnrch 1970. iii + 152 p.• 72 or th.· !i>tn'am for spawning and nurser)" habitat. Cap'~ Fear Rh'cr, N.C.. ri~~ .• :\7 tahles. 1962-66. R)' Paul fl. Nichols and Darrell E. Luuder. Ocwber 1970. iv + 12 p.. 9 ri~s .. 4 tables. 33:1. Het:ommendt'd ppl.cticcs !or Vf'~st'l ... anitation nnd fish hnndl:r.~. By r:dfot'ar' W. Bowman and Alfn·d Larsen. March ]970. iv + 27 p.• 6 figs. 3S6. Flonl~ng Illborator)' for study of aqual1c organisms ar,d their enViron· Rob-~rl Prf)~ress ml·nt. By GeOlgc R. Snyder. Thl'Odore H. Dlnhm. llnd J. McConnell. .;:is. report or the Rurl'41u of Commercial Fl!lherie!. Center for !\1:ty 1971. iii + ]6 p., 11 figs.• El't1l:irine and Mt-:1hadcn Research. Pesticide field Station, Gulf Freeze. F13.• :-is\·:i.! ~'cal 1%9. R.)· tht" Labormory :itarr. Augu~t 1970. iii + 33 p.• 29 figs .• 12 '.l,bll':;, 361. Regional and other related aspect'S of s!le1lfish commmption - some preliminary rindings from the 1969 Consumer Yanl'l ~urv~y. By Morton M. 33',. The Mr\nern fur st'al. By Ralph C. Ba~·,:::,. Ford Wilhe. and Co Howard ,Miller and Darrel A. Nil5b. June 1971. iv + 18 p.. 19 rigs .. 3 tables. 10 apps. 111.lt7.0. April m70. iii + ]9 p.. 13 ri,l"rs. 362. Research vessels of the Nation..1Marint Fisheries Service. By Robert S. :lJ7 _ Pr()~r;lm or Ilh'ision or F:l"onomic R(,!oil'areh, Bureau flf Comlnt'rcwl \\'nl£. August 1971. iii + 46 p.. 25 filS.. 3 table5. For :sale by the i'i~l1t'ril'~. riscal )'l'ar i%9. By Division of £eonomic Research. April 197f·. iii Superintendent of Document!'i. U.S. Goverr.ment Printing OWee. Washingtcin. + :::1') p.. 12 fiJ,rs .. 7 table5. D.C, 20402.
338. Burcliu flf Commcrdal l-'i5hnics Biolngical Laboratory. Auke Bay. :164. Hisltlry Bnd dev':lopmenl or surr clam harvesting gt~r. By Phillip S. Alaska. By Bureau of Commercial Fisheries. June 1970. 8 p•• 6 rigs. Parkcr. October 1971. i ...· + 15 p.. 16 fogs. For sale by the ~:"perintenl:!ent of Documents. U.S. Government PrinLi~f, Ofjjc~. Washington. D.C. 2040:2. 3;m. .'';alrntoi1 rcsrl1rch at h'c Hnrbor Dill'll. r~' Wesley ~l. Ebcl. April 1970. 0 p.. -1 figs. 365. Processin,:: EASTROPAC STD dala and the eonstrt:c:tion or 'H·rtical temperature .'lnd 5ali~,ity sections lJy computer. E)' Forrp~l R. Mille!" ana, ~O. Haft'.m or Comm('\'cinl Fishl'ries Tet"hnolo/oric:al Laborator,Y. Glr'\lc-estl'r. KcnnC'th A. Blis.'\. F'r:t..ruary )972. lV + 17 p.. 8 figs.• 3 appendix fi~. Fer Ma!'sachuselts, By Burr01:J of Cummercial F'ishl·ril's. Junr 1970. l) p•• II figs. saIl' by the Superintendent of Documents. U.S. Government Printing Offi,·'('. W,.~.. hinmo~I, D.C. 2O·m2. . 3-11. Report of the Bureau of Commercial Fisheries Biological Lahoratory. B\~ilUrOrt. N.C., for the riseal year pndlng June 30. 196e.. By the Lllooratory starr. AU~llst nno. iii + 2.; p.• 11 n~s .. HI t:tbles. 360. Kc)' to field identification of andromous juvenile salmonids in the Patific Northwest. By Robert j, McCrlOneil and George R. Snyder. January 1972, iv + 6 p.. 4, figs. F(lr sal~ Jb,Y the Superinttmdcnt or DoculT,ents. lJ.S. :J-l2, !\('~ort of the Burzau of CommerciO\1 fisheries Biological l ..aboratory, Government Printing Ortice. 'A' ashington. D.C. :W402. St Petersbur~ Beach. Florid:t. risc31 year 19(19. B)' the Laboralllry "tolr. AUJ:uSo! .!'?7v. ili + 22 p., 20 rigs.• 8 tables. 367. Engineering ec:onoM.ic model ror ':ish prolein ('~nr.entration pror.esses. By :J.13. Hep"rt of th~ B\ITC'au uf Commercinl Fisheries Biologicol Lab::tralQry, K. K. Almenas. L. C. Du"lIa. R. C. Ern.t. J. W. Gentry. M. B. H:1Ie. and J. Gal\'elitorl. 'fexlI!i. fiscel year 1969. B)' the Laborator)· starr. August 1970. jii M. M~rchello. October 1972:. iii + 175 p.. 6 figs.• Ii tables. For ~ale by the + 3~ p._ 2S riJ:!.... 9 tables_ Superintendent of Documents. U.S. Government Printing O!fi:e. Washinf,ton, D,C. 20402. :\.t,~. nUrl'a~1 or Commercial Fisheri(>5 Tropical Atluntic BidoF,ical I.ahoralor\' prc.gft~~S in research 1965-69. MiAmi. Florida, By Ann Weeks. (}ctoher 1970. iv 3EloS. Cooperath'e Gulf of Mexic:o estuarine inventory and ~tudy. Florian: + "5 po' 53 fig'" Phase t. an!i1 description. B.\· .f. Kneeland Mr.Nulty. WiIHa:n N. LindaU. Jr.. and James E. Sykes. November 1972. vii + 126 p .• 46 fill'S .• 62 t.abies. For 346. Sporlsman's guidl' to handling. smoldng. and preserving- Great Lakes .sale by the Superintendent of Doculnent.s. U.S. Government Printing O[fice coho salmon. B)' Shraron Dudley. J. T. Graikoski. H. L. Sf'agran. and Pau! M. Washington. D.C. 21)402. • Earl. September 1£170. iii + 28 p., 15 rig!7.. 3€9. Field guitl.e tu the ang1efishe~ (Pomaca.nthidae) in the western Atlantic:. 3-:7.. S;)'nopo;i'i o~ bioll~gical data on Pacific ocean perch. Sebc.stodu alutur. ~)' H~nrJ A. Fedderr.. November 1972. iii + 10 p •• 17 figs., For sale by the ~j' Richard L. MaJor nnd Herhert H. Shippet,. Dcccmbu 197a. m + 38 p.• 31 .?up,~rlRte~dent of DOI.;L1ments. U.S. Government Printing O£fice. Wal\hington. fl~s .• 11 tiihles. D,C. 2010.:, Continued un inside b8~'k c:over.
i- h CONTENTS
Page Introduction , ,...... '" 1 Ecology...... 2 -Collecting methods...... 2 Key to the species of marine Tardigrada of the world ,...... 3 Annotated systematic list of marine Tardigrada 01 the world '...... 21 r.ist of marine Tardigrada reported from the northeastern United States .. ,,.,...... 22 Selected bibliography ,...... 22 Indp.x to "cientific names ,... 24 Acknowledgments , ,...... 25 Coordmator's comment 25
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iii Marine Flora and Fauna of the Northeastern United States. Tardigrada
LELAND W. POLLOCKl
ABSTRACT
The manual includ"s an introduction to the general biology. an illustrated key. an annotat"d systematic list. a seJected bibliography. and an index to the Tardigrada of the marioe coastal areas of the world to a depth of 5,000 m.
INTRODUl:TION Since one-half of these are in monO- or ditypic genera, and two-thirds have been discovp.red since 1950, it is likely thai. The Tardigrada (/uTdus, L. slow; ~lradus. L. step) many more species will be described in t.he fut.ure. comprise a phylum of microscopic metazoa (usually less than Marine tardigrades rarely exceed 0.5 mm in length and 1 mOl in length) of uncertain phylogenetic placement. Con are all similar in general body plan Wig. O. They possess as siderations of their status have been based on 1) their many as 11 cephalic appendagcuticle; and 10) the occurrence of eutely or cell constancy especially posterior-lateral cirrus (e) (Fig. I), can he of (although this recently has been disputed, Bertolani 1970). taxonomic significance. In the order Eutardigrada. the 1IIost 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 cephalic characters listed above; in addition. their "ladder-type" appendages ventral nervour. sytem recalls the annelid-arthropod line. \~~ Tardigradan embryology however apparently includes a total but irregular cleavar.e pattern and enterocol'lous / ( fl.'~toe & claw formation of a series of coelomic pouches, of which only the cusp gonocoel is retained in the adult. While this pattern of ,,, I development is unlik.. any other known group, it is most ala-~I ./~toe similar to that of the deuterostomous illvertf:brat"s. Lack of clarity r'cr-arding their phylogenetic affinities suggests that the Tllrdigrad;J. are far removed from their ~ / nearest phyletic n:!ighbor. Apparently they are an old group claw somat iC_""~i-"""-'--7 which has become highly ,:penalized for life in peculiar spine habitats. such as the water filns surrounding lower plants leg spine and lining interstitial space',; between grains of sand. Morphological diversity aP.1or,g marine tardigrades attests ."..--+--gonopore to their age. On the other ha~ld, the comparative uniformity cirrus e---'9' --anus in appearance and simplicit.y in morphological characters of -\- caudal freshwater forms supports the hypothesis that marine appendage tardigrades are primitive. There arc 43 described species of marine tardigradas ineluded in 17 genem. Most ure members of the interstitial meiofauna of sandy sediments. Figure I.-Diagrammatic iUufl'tration of a composite mm;)p. tardigrade. CeJlhalic appendages: B• lateral cephalic drrus; c! . claV'"<.ii ee - external cirrus~ I Department of Z<>Jlogy. Drew Universil)', Madison. NJ 07940. cephalic ic . intt'rnal cephalic cirrus; me - mediaL' cephalic cirrus. number and relative sizes of placoids or rod-shaped (occurring somewhat deeper in beaches under the influenl~p concretions within t.h~ bulbous muscular pharynx can be of heavy surf), The abundant BatiUipes dominates mid-beach diagnostic. sand while other Batillipc/1 often are relegated to more Members of at least six genera (Arckechiniseus. landward or seaward locations. A comparatively denser BatiUipes, Coronarctu,;, Echiniscoides. ParastygarctlLS. and concentration of Stygarctus often occurs deeper within the Stygarclus) display sexual dimorphism in the shape and beach approaching the deepest sediments undergoing tidal location of gonoporcs. Females possess rosette gonoporcs drainage of interstitial water. located mid-ventrally at considerable distance anterior to Less frequently. marine tardigrades are reported from the anus. In males. the gonopore is circular or tubular. other habitats. Sublittoral specimens have been collected to mid-ventral and only slightly anterior to the anus. In a depth of 4,700 m (Renaud-Mornant 1974). Their members of the genus Hal£chiniscus. dimorphism is comparative scarcity in most deepwater surveys suggests exhibited in the relation of the length of clavae relative to either that Tardigrada are less successful here than are the lateral cirri. In males. clavae are longer than lateral many other meiofaunal groups or that sampling and/or cirri. while the opposite is true of fem.des. Sex determi observational techniques commonly used fail to include stIch nation in other marine tardigrades is 'based on presence of small members of the meiofauna. Certain Tardigrada occur mature gametes in the gonad or on the fact that males on seaweed ranging from intertidal Enteromorpha and possess two vasa defer'~ntia while females have a single Lichinia to offshore SaTga&sum. Among the several oviduct. Tardigrada reportedly living ectocommensally with various Tardigrada develop di:-<;etly. Excepting their dispropor other marineinvert£obrates. Tetrakentron synaptae Cuenoi. tionately longer cephalic appendages and their reduced 1892 alone possesses obvious morphological adaptations to number of claws per leg. juveniles resemble miniatuJ'e such a life-style and has been found exclusively in sllch a adults. Growth in Tardigrada is accomplished throu!~h relationship_ periodic molting of all cuticular structures. including the linings of the foregut and hindgut. Apparently internal fl'Jid COLLECTING METHODS pressure is reduced enough t.o permit defecation. ovipl)si tion, and sperm penetration only during an intcrmolt period. A complete discussion of techniques for working with Other aspects of the morphology and anatomy of marine int.erstitial meiofauna general1y and marine Tardigrllda tardigrarles lie beyond the scope of this presentation. specifically may be found in H ·t:,,'ts and Gray (1971). Interested readers are referred to monographs by Marcus Quantitative extraction of tardigrades from sand reGnires (1936). Rudescu 119641. and ftamazzotti (1972). and to a rigorous procedures since most species are strongly recent review by Renaud-Mornant and Polloek (1971). thigmotactic and vigorously resist dislodgment. Anestheti zation by flooding a small sand sample with 3.5% MgCI 2 may ECOLOGY be effective for removing Tardigrada from sediments gathereo in areas of low to moderate wave activity; In recent years. ecological studies of marine Tardigrada however. this technique is not effective quantitatively on ha ve focused largely on thos'! Hving interstitially among samples from "high-energy" beaches (Gray and Rieger grains of sand \Renaud.Debyser 1959~, Schmidt 19'58, 1969; 1971). Soaking small quantities of sand (e.g., 10 ..m] or less) Pollock 1970c; Lindgren 1971). Tardigrades are found in 10 times that volume of 3.5% ethanol is more effective for, throughout portions of intertidal beaches which undergo anesthetization. This can be fohClwed by three or more periodic drainage and replenishment of interst.itial water. rinses of seawater to provide revived and apparently Most interstitial meiofauJla., includingt.llrnier.tl",~_ are unhu:-;n"d Tai'uigraLia qua.ntitatlvely. absent or uncommon in beaches of fine sand (mean grain Marine Tardigrada can be preserved well in either 5·7% diameter less than 300 filn) and in beaehes of larger grain neutralized Formalin or in 70% ethanol. McGinty and size but where finE' silt and debris clog pore space~ :md Higgins (1968) described a widely used techniql,;(' for mount reftrict circulation. ing marine tardigrades. Specimens preserved in 7% Tardigrada occupy specifiC ponions of littoral beaches Formalin are transferred to a 1: 10 glycerin-Formalin creating patterns of zonation both horizontally along the solution which then is allowed to evaporate ~o glycerin (a beach surface and vertically within the sediment. A "typical glycerin-alcohol solution works for specimens preserv&d in pattern" of species cumposition and distribution on a single alcohol). Tardigrad~s prepared by this technique can be beach becomes evident from studies of temperate, quartz mounted in glycerine. glycerine jelly, or Hoyer's medium. sand beaches. An abundant species and from one to several Phase contrast microscopy is necessary for fine observa less ~,ommon :;pecies of BatiUipes occupy superficial sand tions. especially if Hoyer'S medium is used.
2 KEY TO T~iE SPECIES OF MARINE TARDIGRADA OF THE WORLD
The follow jng key is designed for the artificial separation of marine tardigrades. Morphological characters are utilized for easy identification and are noi intended to fully describe the animals. While examination of living animals at high power or oil immersion ;s imperative for complete and accl'l"ale descriptions. specimens fixed in 10% Formalin or 70% alcohol are usually recognizable. The illustrations are variously modified from origin~.l illustrations or descriptions. Important distinguishing features are indicated on each figure by short pointer lines.
Legs terminate in claws which attach directly. or if on toes are longer than toes . 2
Legs terminate in toes without claws or with claws shorter than toes __ . __ . 12
2 (1) Central two claws on each leg bear hairlike extensions , '" , ., , ,...... 3
2 It j Claws without hairlike extensions , .. ,, ,...... 6
3 (2) Caudal spikes absent: anterior margin of head deeply sculptured ,,,.. .. -1 3 (2) Caudal spike present: anterior margin of head much less deeply sculptured ,_.. 5
4 13) Lateral extensions of dorsal plates end in two points .. , , , ., .. " Parastllgarctlts higgiJl.'1'
4 13) Lateral extensions of dorsal plates end in single point Parastygarrt.lls sterreri
3 5 ~3) Somatic spines on mid-posterior border of somatic plate II present; 2 cusps ollly along margin of cephalic plate Stygarctus bradYPlIs
5 (3) Somatic spines on somatic plate II absent; 4 cusps along malgin of cephalic plate Siygarct?Ls gTl4nulatu.~
6 (2) Cephalic appendages absent 7
6 (2) Cephalic appendages present " 9
7 (6) First phar~'ngeal macroplacoid longer than second; inner diam- eter of pharyngeal tube 3jlm 8
7 (6) First macroplacoid shorter than second; inner diameter of pharyngeal tube 1.5 !-1m ...... , " Hypsibius stenostomus
4 B m Second macroplacoid shorter than third Hypsibius appelloefi
B (7) Second macroplacoid as long as or longer than third Hypsibills gp.ddesi
9 (61 More than [our claws (usually 5-11) on each leg; distinct dorsal cuticular plates absent Echiniscoides sigismlllldi
9 (6) Four claws or less per leg ...... 10
5 ' f\;:.':.•.:...... : .•.:.... ~~i' "1 .. \~'?
10 (9) Four claws per leg pres~nt OJ. J.nterior three pairs but only three claws pel leg on posterior !Jail'; distinct dor- sal plates absent; m"ji,m cirrus absent ... .4nisonyches dirlkitlius
10 (9) Four claws per leg pr(:senlon all legs __...... 11
11 110) Median cirrus present; distinct dorsal cuticular plates absent .. " CorOl!!1rclus tcneUus
11 (.fO) Medi~n cirrus absen!; dor~al cuticular pla:~s present Echiniscus rE. (J,1Hyjrispinoslis is the only species reported from marine environment.)
6 12 (1) Four La six toes without claws on ea~h leg...... 13
12 (1) Four toes with claws on each leg '" 27
13 (12) Toes end in disc expansions "" " J4
13 (12) Toes end in narrow lobate expansion;; 26
14 (13) Projecting caudal appel~dage absent 15 14 (13) Projecting caudal appendage present ," 16
15 (14) Caudal p.nd swollen cephalic append- ages long Biitillipes tubernatl:~
15 (14) Caudal end relatively flat; cephalic appendages short BatWipes acuudatll.'i
7 16 (14) Caudal appendage basically a single spike 17
16 (14) Caudal apilendage more than one spike...... ,... 23
17 (16) Caudal spike terminates in a membranous bag Balillipes bul/llCGudallls
17 (16) Cauda! spike blunt or slJarp tipped ' .... , ...... •...... 18
18 (17) Distinet eonstrieU"n{s) present on elavae '...... 19
18 (17) Clava uniform in width. constrictions ab~ent ,.,...... 21
19 (18) Single constriction present on clave .. ,...... , 20
19 (18) More tban one constriction present on clavaE' Batillipcs a1l111l/alu",
3 20 (19) Caudal spike from two-lobed base ... .Batillipes gilmal'1!71i
20 (19) Caudal spille from single-lobed base , Batillipes pemmld
21 (18) Leg spines on hindmost legs short; leg spines present on anterior three pairs of legs. '., "., Batilhpes 1IIirus
21 (18) Legspines on hindmost legs long; spines absent on anterior three pairs of legs...... _ 22
9 22 (21) Caudal appendage a long, slender, sharp-tipped spike .. Batillipes canwnensi.~
22 (21) Caudal appendage, a short, thick. blunt-tipped spike Batillipes s':mili"
23 (16) Caudal appendage t",o·spiked ...... BatilJipes dicrocercus
23 (16) Caudal appendage bears three or more spikes , 24
10 24 (23) Caudal allpendage bears three spikes only . .. 25
24 (231 Caudal appendage bears more than three spikes; central spike longest Bu.tillip'Js phreatieus
25 (24) Spikes of equal length Batillipes littoralis
25 (24) Lateral spikes shorter than central spike Batillipes friaufi
11 23 (lSi Papilla present on fourth pair of legs; clava slightly constricted Orzeliscus belopus
26 (1S) Papilla absent on fourth pair of legs; clava uniform in wirIth Orzeliscus septentrionalis
~~.
f.·•.··· .\~ ". ,'" .. ~.;
27 (12) Middle two toes on each foot much longer thaa outer two toes Archechiniscus marc'i
27 (12) Toes approximately equal in length , 28
12 28 (27) Each claw has more than two exposed points 29
28 (27) Each claw has one CliO two exposed points 30
29 (28) Body nattened dorsoventrally: cephalic appendages reduced; ectocomm~nsu: ..
• 0 • 0 •••••••••••••••• 0 ••••• Tetrakentron synaptae
29 (28) Bolly cylindrical. 0 • 0 •••••••••••••••••••••••••••••••••••••••••••••••••••••••••••••••••••••••••••••••••• 30
30 l29) Earh claw has three exposed points ...... 31
30 (29) Each claw has four exposed points Bathyechinisclts tetrollYx
13 31 (30) Cirri (~and clavae both pre:lcnt 32
31 (80) Cirri c and clavae both absent ...... Styraconyx hapwceros
32 (81) Clavae long (30 ",m); eyes present ,Styraconyx pau!lle
14 32 (81) Clavae short (13 I'm); eyes absent Styraconyx sargo,s;
33 (28) Body covered dorsally by tubules and gelati- nous coating Actinarctus doryphorus
33 (28) Body smooth. lacking tubules.and gelatinclUs coating , .
34 (33) Cuticle rorms two cauda! projections (brancbed or unbranched) extending more than bouy length " 35 34 (33) Caudal projections less than body length or absent...... ,...... 36
15 35 (34) Caudal projections branchtld Tanarctus arborspinosu.~
/
35 (34) Caudal projections 'mbranched Tanarctus lauricus
16 36 (3~) Clava bulb·shaped; less than 0.5 length of lateral cirrus;
eyes present .. 0 •• 00 0 • 0 •• 0 0 0 • 0 0 • 0 o' •• 0 0 • oPleoeola limnoriae
36 (8~) Clava thin. more than 0.5 length oflateral cirrus; eyes
absent 0.0000.0. 0 0 0 0 0 0 0 0.0 ••••••• 00.0 ••• 0 ••••• 0 ••• 0 •• 37
37 (86) All claws crescent shaped without "anchor spikes"
o • 0 0 0 ••• 0.0000.0 ••• 0. 0 0 o' • 0 0 0 0 0 0 .000. 0 0 o •••••••••• 40
37 (36) Crescent shaped claws on middle two toes of each foot with
accessory "anchor spike" embedded in toe 0000•0•0•0•00•00 ••• 0 38
38 (87) Five pl.ripheral alae or membranous sheets present
00 ••••• 0.00 •• 00. 0.0000. 00.0000. 0 0 0.0 •• 00 •••• 0 ••••• 39
38 (.?7) Three peripheral ala~ present; caudal ala smooth, ~!'·.c;"ided; lateral alae undivi<:.:;d .0. 0 0 • 0 o FlQraretus salvati
17
.. ;~ .. ~- .. 39 (38) Caudal ala slightl~· sculptured. not divided; lateral alae divided Florarc/'ls antillensis
39 (.1'8) Caudal ala deeply sculptured and m~arly divided; lateral alae divided Florarclll.~ hcimi
18 40 (97) Peripherai alae p~esent Halechiniscusinlermedins
40 (37) Peripheral alae abt:ent ., , , 41
41 (40) Caudal spike prominent Hauxkiniscus remanei
41 'i40) Caudal s~ike absent . . , "'. 42
19 42 (~1) Cirri" present in addition to three pairs cf somatic spines Halechinisclls guileii
42 (~1) Cirri e present but additional somatic spines absent...... 43
43 (42) Enlarged papilla on shank ofhindmostlegs; body outline smooth " Hakchiniscus perfeel u.~
43 (-..2) Simple leg spine present on shank of :lindmost legs; body with slight sequential ridges ., .Halechiniscus 3ubterraneus
20 ANNOTATED SYSTEMATIC LIST OF Mornam 1961); Renaud·Mornant 1967b). Interstitial in MARINE TARDIGRADA OF coralline sand. Souther!l Pacific. THE WORLD Tana'rclus arborspinos~: Lindgren 1971. Interstitial, inter tidal. Western Atlantic. The arrangement of the following list is based on the Tanarctus tauricus Renaud-Debyser 1959b. Interstitial. classification suggested by RamazzoW (1972). Original intertidal. Western Atlantic. descriptions of all speci::s are available through references in the bibliography. Where subsequent descriptions are useful, FAMILY BAT1LLIPEDJDAE they are cited parenthetically. Finally, the most typicai habitat and geographical range is indicated for each species. Bati/lipel' acaudatus P"IIL'Ck 1971. Interstitial. sandy inter· tidal. Northestern Atl"ntic. Batillipes annulatus DeZio 1962. Interstitial. sandy inter Order HETEROTARDIGRADA tidal. Mediterranean_ Suborder Arthrotardigrada BatiUipes bulJ.u,audtLtus McGinty and Higgins 1968 (Pollock FAMILY HALECHINISCIDAE 1970a). Interstitial, sandy interti.jay. Northwestern and western Atlantic. Ha1echinisr:us guiteli Richters 1908. Interstilial, sandy BatiUipes carnonensis Fize 1957. lnterstitial, sandy inter· intertidal. English Channel, eastern Atlantic. Mediter tidal. Mediterranean. Indian Ocean. ranean. Black Sea. BatiUipes dicrocerr:us Pollock 1970a. Interstitial. sandy Ha1echinisCUll i1/tumedius Renaud-Mornant 1967b. bter· intertidal. Nortlhwestern Atlantic. stitial, subtidal, coralline sand at 4 m depth, Southern :JatiUipes jriaufi Riggin 1962. Interstitial. sandy intertidal. Pacific. Western Atlantic. Halechiniscus perfeetus Schulz 1955. Interstitial. sandy Batillipes gilmartini McGinty 1969. Interst.itial, sandy intertidal and subtidal at 1'10 m depth. North Sea. Medi intertidal. Eastern Pacific. terranean. southern Pacific. Indian Ocean. B... l-iiiipes littoralis Renaud-Deb,Yser 1Q59a. Sandy inter Halechiniscus remanei Schulz 1955 (McGinty 1969). Inter tidal, interstitial. Ea~tern Atlantic. stitial, sandy intertidal to subtidal at 150 m depth. Medi Batillipes mirus Richters 1909 (Marcus 1927; Rudescu 1964: terranean. eastern Atlal'tic. eastern Pacific. McGinty and Higgins 1968; Pollock 1970a, b). Sandy in Haleehiniscus subterrane' , Renauu-Debyser 1959b. Inter tertidal and shallow subtidal, interstitial. Baltic. North stitial, intertidal in coralline sand. Western Atlantic. Sea, Black Sea. M~di\.er!'anean. Atlantic, Indian Ocean. Pkocola lirnnoriae CantacuZEme 1951. Commersal of isopod ,BatiUipes pennaki Marcus 1946 (DeZio 1962; Pollock 197Ua, Limnet,..;a lignorom and sandy int.ertidal to ~ubtidal at . b). Interstitial, sandy intertidal. Northern and south· 130 m depth. F}rlglish Channel, western Atlant.i~. ern Atlantic, Mediterranean, Indian Ocean. Actinarctus doryphoTUs Schulz 1935 (Grell 1937). Intersti· BatiUipes phreaticus Renaud-Debyser 1959a (Riemann 1966; tial, sandy intertidal to subtidal at 170 m depth. One Pollock 1971). Int.ertidal. interstitial in sand. North· report of possibly ectocommenscl relationship with eastern Atlantic. North Sea. Echinocyamus pusillus (Grell 1937). North Sea, English BatiUipes similis Schulz 1955. Sandy intertidal, interstitial. Channel, eastern Atlantic. Mediterranean. Tetrakcntrull synaptu£ Cuenot 1892. Ectocommensal on [li1.tillipes tubernatis Pollock 1971 IRiemann 1966). Sand\' tentacles of holothurian, Leptosynapta gaUiennei. Eng intertidal, shallow subtidal. North Sea, northeaster~ lish Channel. Atlantic. Styracony;c hapwceros Thulin 1942_ Lichens near high OrzelisCUll bewpus Bois-Rcymond Marcus 1952 (Renaud tidal line. English Channel. Mornant 1967b). Interstitial in sand, intertidal and shal Styracony:r paulae Robotti 1971. Shallow subtidal, epizoic low subCdal. NOl'theastern, western, and southwestern on coral. Atlantic. Orzelisr:u.~ Styraconyx sargassi Thulin 1942. Pelagic 011 algae, septent'rWnalis Schulz 195:Jb. 'Interstitial in especially Sargassum. Western Atlantic, Mediterra sand. int.<'rtidaJ. North Sea. nean, Gulf of Mp.xico, eastern Pacific. BathyechinisC1ls tetr'Jnyx Steiner 1926. Questionable de FAMILY CORONARCTIDAE scription based on single observation from mud at 400 m depth in south Atlantk. Coronarctus tenellus Renaud-Mornant 1974. Abyssal mud. FiJJrardu.:; antilknsis Van del' Land 1968 (Renaud-Mornant Southern Atlantic, Indian C"ean. 1971). Interstitial in coralline s:\nd, 3 m depth. Carib· bean. FAMILY STYGARCTIDAE Fl.orarctus heimi Delamare Deboutteville and Renaud Mornant 1965 (Delamare Debouttevill~ and Renaud Stygarctus bnulypus Schulz 1951 (Renaud·:Mornant and An Mornant 1966; Renaud-Mornant 1967b). Interstitial in selme·Moizan 1969). Interstitial, intertidal. North Sea. corallilll~ sand. Southern Pacific. northwestern and northeastern Atlantic, southern Pacific. Fl.orarctus salvati Delamare Deboutteville and Renaud Stygarctus granulatus Pollock 1970a (Pollock 1970b). In Mornant 1965 (Delamare Debouttevill~ and t~c~aud terstitial. intertidal. Northwestern Atlantic.
21 Pamstygarct\/,s higginsi Renaud-Debyser 1965a (Renaud SELECTED BIBLIOGRAPHY Debyser 1965b;. Renaud-Mormmt 1967a). Interstiiial, BERTOLANI. R. intertidal. Southwestern and northeastern Indian. 1970. Milosi somalich. e costanza cellulare numerica nei Tardi· Pa'rastygarctus sterreri Renaud-Mornant, 1970. Sandy gradi. Rend. Acc. Noz. Lincei. Ser. 8a, 48:739·742_ inte,:stitial. Mediterrv.nean. western Atlantic. BOIS·REYMOND MARCUS, E. 1952. On Soulh American Malacopoda. Bol. Fac. Filos. Cienc. Letras. Univ. Sao Paulo, Ser. Zoo;' 17:189-210. CANTACUZENE. A. Suborder Echiniscoidea 1951. Tardigrade marin nouveau, commensal de Limnor;a ag· norum \Rllhke). C. R. Acad. Sci. Paris 232:1699-1700. FAI\IILY OR.EELLIDAE CUENOT,L. 1892. Tm digrade. In: <::Ommensaux et parasites des Echino· Echiniscoides sigismundi (Schultze 1865) iMarcus 1927: dermes (Il). Rev. BioI. Nord. Fr. 5:16·19. Rudescu 1\164: POllock 1975). Intertidal on green and DELAMARE DEBOUTTEV[LLE, C.. and J. RENAUD-MORNANT. blue-green algae growing on barnacles, piers, seawalls, 1965. Un remarqoahle genre de T.rdigrades des s.bles ,~.rallicns etc. Cosmopolitan. d" Nouvelle·Caledonia. C. it. Acad. Sci. Paris 260:2581·2583. 1966. Un nouveau genre de Tardigrarlos des sables detritiques At'Chechiniscus marciSehulz 1953a (Renaud-Mornant 1967bl cor..lliens de Nouvelle·Calildonip. Cah. Pac. 9:149-158. Interstitial, sandy intertidal to shallow subtidal 0_8 DcZl0. S. m). Southern and eastern Pacific. Caribbean. 1962. Descrizione de BatiIJ.ip•.s annulatlL8. n. sp. c nc~e su Batl!, Anisonyches diah;idiu~ Pollock 1975. Interstitial in coralline lipes pennaki Marcus. nnovo rinvt::nimento nel Med~terraneo sand. Westl'rn Atlantic. eastern Pacific. (Heterotardigradal. Annu. [st. Mus. Zoo\. Univ. Napoli 14:1·7. FIZE. A. 1957. Description d'une espe.e nou""lIe de Tardigrade BatiIJipes FAMILY ECHINISCIDAE carnonenGis n. sp. Bull. Soc. Zool. Fr. 82:430·433. GEDDES. D. C. Echiniscus quadri.~pinosus Richters 1902. A moss water 19GB. A note on the marine tardigrade H'III'Bibius (lsohypBibius) film species found in marine setting only once (Renaud ste1Wstomus (Richters) from the Trams. area. Northern Norwnl. Debyser 1964). Eastern Atlantic. Astarte 33:1-3. GRAY, J. S., and R. M. RIEGER. 1971. A quantitative sludy o[ the meiofauna of an exposed sandy beach, at Robin Hood's Bay. Yorkshire. J. Mar. BioI. Assoc. Order EUTARDIGRADA U.K. 51:1·19. GRELL. K. G. FAMILY MACROBIOTIDAE 1937. Beitrage zur Kenntnis von Actinat
22 1975. Observations on marine Heterotardigrada. including a new RIGGIN. G. T. genustrom the western Atlantic Ocean. Cah. Bio1. Mar. 16:121 1962. Tarciigrada o[ southwest Virginia: with the addition at a 1.32. description of a new marine species from Florida. Teeh. Bull. RAMAZZO'ITI. G. Va. Agric. Exp. Stn. 152:1·145. 1972. 11 phylum Tardigrada. Mem. 1st. lta!. Idrobiol. 28:1·732. lVlBO'IT1, C. 1971. Nuova specie di tr.rdig;-ado marinll: Styraccmyz: paulae RENAc,D·DEBYSER, J. (Heterotardigradal. Ann. Mus. Civ. Slor. Nat. "Gi:oeomo Doria" 1959a. Sur quelques Tnrdigrades du bassin d'Arcachon. Vie 4(199):1·3. Milieu 10:135-146. RUDESCU, L. 1959b. Etudes sur la [aune interstitielle des nes Bahamas. III. 1964. Tardigrada, Fauna Republicii Socialiste Populare Roma· Tardigrades. Vie Milieu 10:296·302. nia, Arthropoda 4(7):1.400. 1964. Note sur 1a loune interstitielle du Bassin d'Arcnchon et de SCHMIDT, P. scription d'un Gastrotriche nouveau_ Cah. Bioi. Mar. 5:111-123. 1968. Die quantitative VP.rt~Uur.g und Populatiunsrlynamik des 1965a. Parastygaretus higginsf n. g., n. sp. Tardigrade marin Mesopsammons am Gezeiten-Sandstrand der Nordseeinsel Syltl. intersliliel de Madagascar. C. R, Acad. Sci. Paris 260:955-957. Faktorengeruge und biologische GIiCl:·erurlg des Lebensraumes. 1965b. Etude sur un Stygarctid€ (Tardigrada) nouveau de Mada· Int. Rev. GesalOten. Hydrobiol. 53:723·779. gascar. Bull. Soc. Zoo1. Fr. 90:31·38. 1969. Die quantitative VerteUung ond Populationsdynamik des RENAUD·MORNANT. J. Mesopsammons am Ge7.eiten·Sandstrand der Nordsee-lnsel Sylt 1967a. Pa'!'fl8tygaretus higginsi Renaud-Debyser, 1965, sur la cote II. Quantitative Verteilung und Populationsdynamik einzelner orientale de Malaisie. Descript',on de la femelle. Bull. Mus. Arten. Int. Revue Gesamten Hl'drobiol. 54:'35-174. Nat!, His!. Nat.. Paris 39:205·208. SCHULTZE, M. 1967b. Tarnigr"ces de \a Baie Saint·Vincent Nouvelle·Caledon!e. 1865. Eahiniscus sigismundi. ein AI·etiscoi.le der Nordsee. Ark. Editiuns de Fo"dat. Singer.Polignae, Paris:l03-119. Millrosc. Anat. 1:428·436. 1970. Parastygaretus sterren n.•p., Tardigrade marin nouveau SCHUl.Z, K de l'Adriatique. Cah. BioI. Mar. 11:355-360. 19~. Actinarctus dOf;}o'p"'Jnrus nov. gen. nov. spec.. ein mel'kwUr· 1971. Tardigrades marins des Be"mudes. Bull. Mus. Nat!. Hist. diger Tardigrad ai!' der Nordsee. Zool. Anz. 111 :285·288. Nat.• Paris 42:1268·1276. 1951. Vber Stygarcim bradypus n. g. n. sp., einen Tardigraden 1974. Une nouvelle famille de Tardigrades marins abyssaux: les aus den: KUstengrundwasser, und seine phylogenetisehe Bedeu· Coronarctidae fam. nov. (Heterotaroigrodal. C. n. Acad. Sci. tung. Kiel. Meeresiorsch. 8:86-97 Paris 278:3087·3090. 1953a. Eine neue Tardignuien·Gattung von der :>azifischen KUste. RENAUD·MOm_ANT, J.• and M.-N. ANSELME-fI'iOIZAN. Zool. Anz. 151:306·310. 1969. Stades larvaires du Tardigrade marin Stygl1.i'etus bradypus 1953b. Orzeliscus septentriDnalis nov. spec., ein neuer mariner Schulz et position systEmatique des Stygaretidae. Bull. Mus. Tard.grad an der deut.chen NordseekUste. Kiel. Mearesforsch. Nat!. Hist. Nat., P~ris 41:883-893. 9:288·292. RENAUD·MORNANT, J.. and L. W. POLLOCK. 1955. Studien a..'1 marinen Tardigraden. Kiel. Meeresforsch. 1P-71. A review of the sl'stematics and ecology o[ marine Tardi· 11:74-79. grada. Smithson. Cont.-ib. Zool. 76:109·117. STEINER, G. 1926. Bathyeelliniscus tetronyxn.g. n.sp. Ein neuer marin...rTar RICHTERS. F. digrade. Disch. SUdpolar.Exped. 18:478·481. 1902. Beitrage zur Kenntnis der fauna der Umgebund von Frank· furt a. M. Ber. Seckenb. Naturforsch. Ges. 33(2):1-21. THULIN. G. 1942. Ein ""uer mariner Tardigrad. Medd. Goeteb. Mus. Zool. 1908. Marin" Tardigraden. Zoo\. Anz. 33:77-85. 99:1·10. 1909. Tradigrader,.Studien. Ber. Senckenb. N,turforseh. Ges. UHl.IG, G. 40(2):28·44. 1968. Quz.ntilative methods in the study of interstitial fauna. RIEMANN, F. Trans. Am. Microsc. Soc. 87:226·232. 1966. Die interstitieHe Fauna im Elbe·Aestuar. Verbrei!","!> 'lDd VAN DER LAND. J. Systematik. Arch. Hydrobiol.. Suppl. 31, 279 p. (see p. 223·225 1988. FWrtlTetus antiUemis. a new tardigrade from the cor'll sand and Fig. 57). o[Cura~ao. Stud. Faua~ Cura~ao Caribb. lsi. 25:140·146.
23 INDEX TO SCIENTIFIC NAMES
A ctinarctus salvati 00 •• 0 .,. _ ••• 0 ••••••• o. 0 0" • 0 0 0 0 0 0" o. 0 17,21
doryphorus . 0 0 •• 0 , 0 0 •• 0 • 0 0 • 0 '00 0 •• 0 0 0 • 0 0 • 0 •• o. 15, 21 Halechiniscidae .... 0 0 •• 0 0 •• 0 0 •••• 0•000•0•0•000, 21
A nisonyches HalecMniscus 0 0 0 0 • 0 0 ••• 0 0 •• 0 0 ••• 0 0 0 •••• 0 •••• 0 •••• 0 2
diakidius 0 • 0 00' •• 0 ••• 00.0.0000.000 ••••••• 0 0 0 .0' 6,22 guiteli 0 ••• 0 0 0 •• 0 •• 0 •••• 0 •• 0 • 0 0 0.·. 0 •• 0 •• 0 •• ' 20, 21
Archechiniseus .00000 ••• 0 • 0 •• 0 • 0 0 0 • 0 0 •• 0 • 0 ••• 0 0 0" 2 intermedius 0" 0 ••• 0 •• 0 • 0 0 • 0 • 0 •• 0 • 0 0 0 0 • 0 •• 0 •• 0 19,21
marci 0 0 • 0 0 0 • 0 •••• 0 • 0 0 0 0 ••• 0 0 •••• 0 •••• 0 • 0 •• o. 12, 22 per/edits 0 0 0 ...... 0 •• 0 ..... 0 ... 0 0 0 .... 0 .... 0 0 20. 21
Arthrotardigrada 0 0 •• 0 • 0 •• 0 •••• 0 ••• 0 0 • 0 • 0 0 • 0 •• 0 0 0 0 21 Temanei " 00•0000•0•0 •• 0 • 0 0 ••• 0 •••• 0 0 0 •••• 0 0 19,21 ~1 Bathyechiniscus subterranCd-s > •••••••••••• •• 20.
tetronyx 00•000•0•00•0•00 ••• 0 • 0 0 • 0 •• 0 •••• 0 •• 0 13, 21 Heterotardigraoa 0 0 •• 0 • 0 0 0 0 •• 0 • 0 0 ••• 0 0 0 • ' • 0 0 • •• 21
Batillipedidae .... 0 • 0 • 0 •• 0 • 0 0 0 0 0 0 ••• 0 0 • 0 ••••• 0 •• 0 o. 21 Hypsibius
Batillipes 0000•00•00•00 ••••• 0 •• 0•000•0•00•0•0•00•0•0 2 appeUoefio. 0 •••• 00 000.000.0. 0 •• 0 ••• 0 • 0 •• 00. 00 5,22
acaudatus 0 0 0 0 0 0 0 0 •• 0 • 0 •• 0 0 0 • 0 • 0 0 • 0 •• 0 ••• 0 0 • 0 0 7, 21 geddesi . 0 0 0 0 0 0 ••• 0 0 •• 0 0 • 0 •••• 0 0 0 0 •• 0 •••• 0 • 0 0 5, 22
annumtus 0 0 0 • 0 ••• 0 •• 0 • 0 • 0 0 • 0 •••• 0 0 0 0 ••• 0 • 0 • o' 8, 21 stenostomus ..... 0 0 0 •• 0 • 0 0 • 0 0 .. 000 0 0 0 • 0 0 •• 0 •• 0 0 4,22
bullacaudatus .... 0 0 ••• 0 0 0 0 0 0 0 •••• 0 • 0 0 • 0 • 0 o. 8, 21, 22 Macrobiotidae .' .. 0 0 0 • 0 •• 0 • ' ••• 0 0 •• 00•0•00•000000• 22
carnonensis 0 0 •••• 0 • 0 •• 0 0 •• 0 0 0 0 •• 0 0 •• 0 0 • 0 0 • 10, 21 Oreellidae .. 0 0 0 • 0 0 ••• 0 0 •••••••• 0 0 •• 0 •••••••• 0 0 0 0 0 • 22
dicrocercus ..... 0 0 •• 0.00. 0 0 0 0 ••• 0 •• 0 • 0 0 • 0 " 10,21,22 Oneliseus
frin.ufi 0.·· •• 0 •••••••• 0 0 ••• 0 • 0 0 •• 0 •• 0 • 0 0 •• 0 o. 11, 21 belopus 0000000•000•0 •• 0 • 0 •• , 0 0 • 12, 21
gUmartini 0 • 0 •••••••• 0 • 0 0 ••••••••• 0 • 0 0 •••••• 0 o. 9, 21 septent'rionalis 0 0 0 0 •• 0 0 • 0 ••• 0 o ••• 0 0 •••••• 0 0 0 0 • 12, 21
littoralis ., 0 ••••• 0 ••• 0 0 ••• 0 • 0 0 0 0 •• 0 0 0 • 0 0 0 •• 0 • 0 11, 21 Parastygarctus o. 0 0 •• 0 • 0 • 0 •• 0 0 •••• 0 •••••••• 0 0 0 •• 0 • • 2
mira~ 0 ••••• 0 0 •• 00•00•00•000 ••• 0 0 ••• 0 • 0 0 o •••• 0 0 • •• • •• 0 •• 0 0 •• 0 •• o ••••• 0 • 0 •• 9,21,22 higginsi .... S, 22
pennaki. 0 0 0 •• 0 0 0 0 •••• 0 0 0 ••• 0 0 0 0 0 •• 0 _ • • 9,21,22 sterreri , .. 0 0 0 •• 0 •• 0 0 •• 0 •• 0 • 0 0 •• 0 • 0 • 0 0 0 0 ••• 3, 22
phreaticus 0 ••• 0 0 •• 0 0 0 0 •••• 0 0 •• 0 •• 00•0•0000•0 •• oIl. 2 i Pleocola
si'l1u:lis . 00•0•0•000•0 •• 0 • 0 0 0 ••• 000000•0000•0• o' 10, 21 limnorin.c. 0 0 0 ' 0 0 •••• 0 •• 0 • 0 • o •• 0 •••••• 0 •••• 0 •• 0 17,21
t'Lbernatis . 0 ••• 0 • 0 0 •• 0 0 • 0 0 ••• 0 •• 0•00000•0•0 •• 0 0 7, 21 Stygarctidae 0 ••• 0 • 0 •••••••••••• 0 •••• 0 •••••• 0 • 0 0 • 0 0 21
GJronarctidae 0 •• 0 0 •• 0 0 ••••• 0 ••••• 0 ••••• 0 •• 0 • 0 0 0 •• 21 Stygarctus 0 • 0 0 0 0 ••• , 0 • 0 0 0 0 0 • 0 •••• 0 ••••••• 0 •• , •• 0 0 • 2
CoronaTetus 0 •••• 0 0 0 0 • 0 •• 0 0 0 0 • 0 • 0 0 •••••• 0 •••• 0 • 0 0 • 0 2 bradYPllS 0 • 0 0 00 •• 0 0 •• 00 ••• , •• 0 •• 0 •• 0 • 0 0 0 • 000 4,21,22
teneUus 0 ••• 0 0 • 0 • 0 0 0 0 •• 0 • 0 0 ••• 0 0 ••• 0 •• 0 •••• 0 ., 6, 21 granulatus . 0 0 0 0 • 0 0 ••• 0 0 •• o. 000000000000 •• " 4,21,22
Ecbiniscidae .. 0 0 0 • 0 0 • 0 •• o •• 0 0 0 0 0 0 ••• 0. 0 • 0 ••• 0 0 0 0 0 0 22 Styraconyx
Echiniscoidea 0 • 0 0 0 • 0 ••• 0 0 • 0 0 0 •• 0 • 0 •••• 0 0 • 0 • 0 0 • 0 • •• 22 haploceTos 00 " 0•0 000•0•0 •• 0 • 0 • 0 0 • 0 , o. 14, 21
Echiniscoides . 0 0 ••• 0 • 0 • 0 ••••• 0 0 • 0 ••• 0 ••• 0 0 0 0 0 • 0 • 0 o. 2 paulae 0 •••• 0 0 • 0 • 0 • 0 •• 000•00•00•0 •••• 0 0 •• 14, 21
sigismundi . 0 •• 0 •••• 0 • 0 0 0 0 ••• 0000 ••• 0 • 0 0 •• 0.00 5,22 sarga.ssi 0 ••• 0 0 0 0 •••••• 0 •• 0 • 0 0 • 0 • 0 0 •••• 0 15, 21 Tanar~tus Echiniseus o' 0•0•0•00000 ••• 0 • 0 •••••• 0 0 • 0 • 0 • 0 •• 0 0 0 o. 6
quadri.spinosus .... 0 ••• 0 • 0 ••••• 0 • 0 • 0 •• 0 0 ••• 0 0 • 0 6, 22 arborspinosus 0 0 0 0 0 • 0 ••• 0 0 • 0 • 0 ••••••••••• o. 16,21
0 ••• 0 0 0 • 0 ••••• 0 0 •• 0 ••••• 0 ••• 0 ••• 0 o' Eutardigrada 2, 22 tauricus 0 0 • 0 0 o. 0 ••• 0 0 0 0 0 0 0 •••• 0 • 0 0 0 0 •• 0 •• 00 •• 0 16,21 Florarctus Tetrakentrun synap~'2e antilJ.ensis 0 0 • 0 ••••• 0 0 0 0 • 0 • 0 •• 0 ••• 0 0 0 • 0 0 18, 21 ... 0 0 • 0 ••• 0 • o. 0 • 0 •••• 000 0 0 0 0 0 0 •• 00 0 2, 13, 21 ~l heimi 0•0•0•0•0000 ••• 0 •• 0 •• 0 0 • 0 , •• 0 •••••• 0 0 0 0 18,
24 ACKNOWLEDGMENTS
l'r"paratifln of the "Marin" Flora and Fauna of the North"ast"rIl Robert T. Wilce, Department of Botuny, University l'niled States" is being coordinated by the following Board: of Massachusetts, Amherst, Mas,.
Coordin:llor: Melbourne R. Carriker, Marine Studies Center, College of Marine Studies, University of Dela· Thl' Bm.rd
COORDINATOR'S COMMENTS
Puhli{'ation of the "Mar-jnt.' Flor~l .md Fauna of thl' Northe:l~tern Npw Hampshire in 1969 for work on tl:e hi(J'c.gy of intf'l'~jrial Unill·d Stat<~s" is most tinw1y in view of thl~ growinK univursal 'I'ardigTada. Both this docl'Jral resl.·are" arld lhl' mz.jorit;,-· of t'lIlphasis Oll l'nvironmenlal work :md thp ur!-:'l'nl need for mOTt' SUh~('qUllnl [>ostdodoral rm';('ar(~h was ('onduetNI :,11 m.soeiation wit h pn'l'is(' and complete identifi('ation of ('oastal or~:lnisms than has t.hp Systt'maties-)<:eolog-y :·;)rol!r:l.m at the \1arine Biolo~ic:1.1 hl'('n 'lvailahlp. It is mandatory, wherever pnssihll', that organisms Lahorntory in \Vouch; Hole. In 1972. he mov.ed to his pi'esent IW if!enlified act'uratel,)' to species, At'{'ural(' :wicillifit· name::. unluck po:-;ilion a~ a meml)l'f of thv ;".Oolog)' fuculty at Dl~t:W University. ttw g-rt'al Cjuanlitie!" of hiohlJ{ical information slon·d in lihrilrit, .... ~'ladi~(ln. N ..]. wiU're his ~llJdi~s on a~pl·l.~ts uf ~_he syslcm:ltil's and ohviittl· duplifulion of rl',search lIin'ady done, ..tnti make pos.sihle I'I'olog-y of marine Tardigr.ula eontinut>, pn·didiull of altrihule."i of org-anisms t.hat havi' lWt~n inadcquiltt'ly 1\'1anuals art' availahle for pun~h"lse from ita' Superintendent of SllHtif'd. Oncumcnts. U.S Gov('rnment l'rill~jnv. Offi('l:. \Vashin1-;wn. D,C. Leland W. l'ollOl:k was awarded his Ph.D. from the Universit)'llf 20402. The manuals ~o far publishcc.\ in the sr:ries are Iisll'o helow.
COOE" !):\VID G.. ;!~d Rl\tJPH O. THUNKHl.lRST. Marim' nora and [tiuna of thl' nOl'tlwastl'rn lJnill'et Stah·s. Ann.'lida: Oligo(·liitC·!;,. BOHH01{, AHTlIlIH C. ~larinl' flol'a and fallna of thl- northl'asterr. lInill'd Stall',. Protozoa: Ciliopho!'a. 1I10l 11., r:DWIN T. Marine flora and fauna of thl' northeastern United SUites. lIi!(hl'r plants of Ihe ",arilll' fringe. ~kCLOSKr;Y. I.AWRENCE H. M""ine nora "nd fauna of til<' nlJrtlll'asll'rn United Stall'S. Py('no!:onida. ~lANNING. RAYMOND B. Marinl' nunl and fauna of Lhe' n(lrtheast~'rn linill'O :-:;lates. CrUsLI('t'a: S!omatopt,d;l. WILLIAMS. AUSTIN B, Marine flor" and fauna of the northeastern Unill'd Statl's. Crllstal'l'a: De('apod". POLLOCK. LELAND W. Marine flora "nd fauna of the northe"slern lInit,·d States. T:lCdi!:eada.
25
'0 u.s. GC'/£RNMENT PRINTING OFFiCE: 1976-697·069 r78 REGION 10 3'm. l:ullccLing and processing data on Ii:.. i1 eggs Bf1d larvae in th~ Califo"nia ~'81. Fishery publications. calendar year 1967: Li!lt.'i and indexes. OJ Lee e. Gurrent region. Dr Da"id Kramer. Mary J. Killin. Elizabeth G. Steven:;. Thorson and Ma.ry Ellen Engett. Jul}' 1913. iv + 22 p•• 1 fig. For sale b)/ th,e ,fames R. Tlmulkin. and Jaml'5 R. 7.v.. ~ir('1. Nt,vcl:lher 197~. iv -;- 38 p., 38 Superintendent or Documents. 'J.S. Govemmt'nt Printin,!:: orne.:. Wasltingto:1, figs .• 2 lllhit's. For salt' by the Superintendent of Document'>. U.S, Govern· n.c. 20402. . ment PrintinR Ofrice. Wl\:-;hington. D.C. ~0402. 38:.!. Fi:.;hery publicIltions. calendar yf03r' 1966: Lists ant inltexes. Ry Mary Elll'n En!Z'etl and Lee C. Thor~on. July 1973, iv 19 p., 1 fig. For sale b)' 311. Ocean fisht!ry management: Discussions and research. D)' Adam A. + the S'Jperinll'ndl'nt of DDcuml·nL... U.S. Gov~rnment Printing Orrice. W89hing· Sokoloski leditorl. 117 pap~rs, 24 authors.) April 1913. vi 173 ;.... 38 figs .. + ton. D,C, 20402. 32 tables. 7 appendix tablt·s. :~3. Fishpr)' publi{,litions. calendar yrar 1965: Lists and indexes. By Lee C. 312. Fisher)' Plj~liclltions. calendar year 1971: Lbt~ and im::!ut"'i. Ry Thoma..s Thon"m and Mar)' EUl'n l':nge\t. Jul)' i913. iv + Ii! p., 1 fig. For salt: b)' the A. Manar. O('l,;·:,ber 1972. iv + 24 p.• 1 fig. Fer sale by the Superintend('nl of ~upt'rinlenden~ of Documents. U.S. Gov~rnmcnt Printir,g Office. Washing Documenl,s. U.S. Go\o~rnment Printing Qtrice. WashiaKton. D.C. 20402. ton. D.C. 20402. ~.t. Murin.' nora and faunn or tt'll' north~a5tcrn United Stat('s. Higher pl"nt... nf tht· marine frinJ:"C'. By Edwin T. Moul. September 1973, iii + 60 p.• 109 figs. For sale by the Superintendpnt or Documenl'l. U.S. Gevern:nent Printing Orrin.'. WashiDJ;1on, p.c. ~'O40:!.
:ns. Nl'W ?olyehaeta from Beaufort. with II k('y to all spe~i(H; Tt!corderi 'rem aS5. Fi.~herJ' publi~'ations, calendar yenr 1972: Lis\.5 and indt:xe~. By Lee C. North Carolin•. lly Joh" H. nay. July 1973. 'iii + 140 p., 18 figs.. 1 ubi•. Thor.;on and Mar)' EUf'n Io..llgett. ~ovember 1913. iv + 23 p., 1 fig. For sale For saIl' h:.· the SUJl~rinlendt'"t of J)ocuments. U.S. uo·.·crnmcnt Printing b)' the ~upcrin~C'ndt~nt of DOCl\mt'h~5, U.S. Govprnment Printing Office. OHice. Wn~hin!'lton. D,C. 20402. W:L"ihinl{tnn. D.C. 20402.
376. Bottom·waler temperatures un the continental shelf, Nova Scotia to :~~~. B~:lr~~r~~~~a R:,dM~tl~:k~J. ~:p~eo=:ee:s:;;~. ~rit+d l~ta;~~-l Pl~~O~~r New J£r~ey. By John H. Colton. Jr. b.nd Ruth R. Stoddard. Jl~:'{, J913. iii + t 55 p.• 15 fins., 12 appl'ndix tables. ror sale hy the Superintendent or sail' b).' the Sup('rintendent of Documt·nls. U.S. Government Printing Office. Dorum('nt~, U.S. Government Prinlinl': QUite. Wa.c;hinJ{1.on. D.C. 20402. I,\'nshinforton. D.C. 20.J02.
377. Fishery publi'·ali(l:ts. cal\~ndar year 1570: Lists and indexes. Dy Mary 387. Marini' nt.rn and faunu of tht' northeastern United States. Crustacea: Ellen Engen and l..e
318-. Mnrint· nora and {nuna ur th£' norlheash'.n Unit__od SlaLes. Protozoa: :iRB. PnK'l'(·dinKs of the firct V.S.-Japan mt~ting on aquaculture at Tokyo. Ciliophora. ")0' Arthur C. Borror. Sept~r:")er 1973. iii + 62 p .• 5 fiR'S. For sale .Jnpan. O('torn..·r IS:1Y. 1~71. William N. Shaw (editorl. (l~ papers. 14 author.'i.l by the Superintendent of Documents. U.S. Government Prinlin~ Orricc, r't'nrunry 197·1. iil + 133 p. For sale by thl!' SUpl'rintel"ldent of Documt.'nts. Wa~hinKlon. D.C. 20402. U.S. GO'{l'rnment PrintinK Orrin-. Wl1shin,.,'1oTI. P.C. 204(Y.!.
379. Fishery publicaticns. calendar yt·.lr 1969: Lists and indexes. B,' Lee C. :!89. Ma.rine nor:l :trId falma or the nnrthetlst('rn United Stntes, Crusl.acea: Thorson and Mary Ellen EngElt. April 1913, iv + 31 p.• 1 fig. For sa~.e by DI"~apoda. 11)' Au ...tin B. WiIIi:lms. April 1974. iii + 50 p .• 111 figs. f~r sale the Supuirllendent of Doc\lme:lt~. U.S. Government Printing Ornec. WashinJ:' b:r thl> f'lJpcrint~ndcnt nl n(Y.'umt:nts. U.S. Guvcrnmcnt Printing ·nrCice. lon. D.C. 20402. Wast.ington. n.c 20402.
380. Fisher~' publictllions, cOlhmdllr yt':lr 1968: List!l and indexes. By Mary :MJ. Fisher)' pullikatil'lns. ('alendar )'ear 1973: Lill;t"i and ind~xes. By \1ary Ellen Engett and Let' C. ']'horson. May 1973. iv + 2-1 p .. 1 fig. For ~ale by Ellen Engell Dnd Lee C. Thar'son. September 1974. iv + 14 p., 1 fi(, For thl' Superinte'ndent of n~i1merll"i. U.S. GmlPrnml'nt Priming QUice. W.:l,ihing· ,,·'.Il· hy tht' Supl'rintendent of I)ocuments, U.S. Government Prinling ·O;ricc. ton. O.e. ~0402. Washinh'1.on. D.C. 20402,