FRESH-WATER INVERTEBRATES OF THE UNITED STATES PROTOZOA TO MOLLUSCA THIRD EDITION ROBERT W. PENNAK Emeritus. resent Gul Biology University of Color Boulder, Colorado WILEY AWiley-Interscience Publication JOHN WILEY & SONS, INC. New York + Chichester - Brisbane + Toronto + SingaporeCopyright © 1989 by John Wiley & Sons, In Reproduction or translation of any part ofthis work Iheyond that permited by Scetion 107 of 108 of the 1976 United States Copyright Act without the permission f the copyright owner is unlawful Requests f Library of Congress Catling in Publication Data: Pennak, Robert W, (Robert William) ited in the United States of America11 TARDIGRADA (Water Bears) Although tardigrades are strictly aquatic animals, they are seldom abundant in the invertebrates. They are not plankton organ: isms; they are only occasionally collected in aquatic mosses and algae, on rooted aquatics, or in the mud and debris of puddles, ponds, and lakes. More typically, active tardigrades are found in the droplets and film of water or angiosperms with a rosette growth form. They also occur in the capillary water between the Sand grains of sandy beaches up t0 2 or 3 mn from the water's edge Mature individuals range from 50 to 1200 yam in length butare usually less than 500 pm long. Many species have a considerable size ariation. The short, stout, cylindrical body, the four pairs of stumpy lateroventral le and a deliberate“ pawing” sort oflocomotion produce a fancied resemblance toa miniature bearlike creature, and the Tardigra been generally called “water bears” since 1869. great majority of species have been reported from lichens, liverworts, and terres trial and fresh-water mosses, All other species have been reported from marine habitats, mostly interstitial Nearly all of what we know about tardigrade biology has been contributed through the efforts of British, investigators. Water beats have been badly erman, Italian, and French neglected in the United States, even though we undoubtedly havea rich fauna, Within th past 20 years, however, there has been in- ¢ des by American investigators. About 90 species have been reported from this country, and this figure 254 includes marine, moss, and. fresh forms, About500 species have been described the world over. General characteristics. Tardigrades are fascinating objects for electron and low microscopy investigations, The body consists of a distinct head and indefinite “body segments.” Each leg is armed with four claws or two pairs of double claws. Crawling and creeping are more or less sluggish, the claws being used for clinging to vegetation and debris. The mouth is anterio al or ventral, and the cloacal aperture, or anus, lies between the bases of the last legs. The cemire surface of the body is covered with a complex, multilayered cuticle secreted by the thin hypodermis, which consists of p subterm| jgonal cells. Depending on the species, the cuticle may be variously thickened and marked; usually there are also various spines, papillae setae, or hairs. Cilia do not occur in this phylum. Only liquid materials are ingested, and the anterior part of the digestive tract has a sucking pharynx and a pair oflong, piercing stylets Sexes are separate, but the great majority of individuals are females. Usually there are four to six ecdyses in the life history A remarkable feature of most aquatic and semiaquatic fresh-water tardigrades is their bility to tide over dry perio sin ashriveled, rounded, anabiotic state. When normal moisture conditions return, the animal resumes its normal appearance and activity Coloration is variously produced by the pigmentation of the cuticle and hypodermis, dissolved materials in the body fluid, reserve food brow relath older pale Th about pierce Oceas smal ronife Th of the smal Fic Schustfood bodies, and the contents ofthe digestive tract. Gray, bluish, yellow brown, reddish, or wn are common colors. Some species are atively dark and opaque, especially among older individuals; others are translucent and pale There are no special circulatory or respira tory systems, Materials diffuse and circulate about easily in the body fluid, and respiration eccurs through the general body surface Feeding, digestive system. Tardigrades aremostly plant feeders, The cellulose wall of moss leaflet cells and algal filaments are pierced with the stylets and the fluid contents aesucked out by pharyngeal pumping action, Occasionally, howeve small metazoans, such as nematodes and tmifers, are used as food. Milnesium tardigradus i sid to be chiefly carnivorous, The periphery of the mouth is stiffened and surrounded by two or three folded and Superimposed rings of cuticle the body fluids of That portion of the digestive system extending from the small mouth cavity to the anterior end of the Fie I.~Scanning electron micrograph of Parech Huser, University of California, Davis, TARDIGRADA 255 ¢sophagus called the buccal apparatus Fig. 5). It consists of the tubular pharynx. th muscular or sucking pharynx, and the stylet mechanism. The tubular pharynx is narrow and sclerotized and extends into the mass of the sucking pharynx. The latter is football shaped, with thick walls composed of radia muscle fibers; its small triquetrous cavity is lined witha thin epithelium. Six longitudinal Tidges or fragmented sclerotized bars give stifiness to the sucking pharynx and provide for insertions of muscle fibers sclerotized pieces in the 5 Theindividual Fynx are called single small posterior piece. called a placoid or comma, The two stylets ieatan angle on either side of the tubular pharynx macroplacoids (Figs. 5,6 and the sharp tips Project into the anterior end ofthe pharyngeal cavity through special lateral slots, or stylet sheat A transverse suppom extends from the tubular pharynx to the basal cach stylet. Several pairs of small muscles extending from the base of the stylets to the tubular pharynx and the sucking pharyns arc capable of withdrawing and protruding the 00, a Semiaquatic species. (Courtesy R. O.tron micrograph of Macrobiot California, Davis, stylets a considerable distance out of the A pair of large salivary glands open into the mouth cavity. They also function in secretion of new stylets and stylet supports molting and for called stylet glands. Some investigators believe that they havean excretory function, especially the Heterotardigrada. A few small unicel lular glands of unknown function occur near the mouth. They are ingrowths of the hypo- dermis. The esophagus ranges from long to very short. Itempties into the capacious stomach, nidgut, which is usually without diverticula but may have slight lateral caeca. The hindgut fectum, is short and has a small cavity. In the Class Eutardigrada itis atrue cloaca, The anus or cloacal aperture is a longitudinal or Digestion in the anterior portion of the stomach is acid; posteriorly it is alkaline Reserve food may be stored in hypodermal cells or in special reserve food bodies which float about in the body cavity. Feces are often released into the old cuticle during molting, especially in Echiniscus Excretion. In the Eutardigrada there are Uree small glands situated at the junction of OF THE UNITED STATES. X150. (Courtesy of R. O, Schuster, University midgut and hindgut, each of which consisis of only three cells. One gland is dorsal elateral. All supposedly have an exere tory function, and perhaps they osmoregulatory. The paired glan monly called Malpighian tubules, but perhaps are more appropriately nephridia ‘etory material is stored in the hypodermal cells, some excretory granules are left within the old exoskeleton at ecdysis, and some are given off at the anterior end when the style mechanism is shed just preceding an ecdysis Musclesystem, Inaddition o the pharynx and the syle muscles, a tardigrade usualy has from about 40 4 140 long, thin body muscles, che number depending on the species Each such muscle is merely a single Abril cell with one nucleus, or several suc body wall Fig. 7). Contractions ofthe dora d ventral longitudinal muscles bring about shortening or curvature of te body Each legis moved byasct ofmuscles origina ing on the dorsal and vental body wall and inserting near hep ofthe leg. The body wall is devoid of cirealar muscle fers, but the Muscles extending from the body wal to the aslTARDIGRADA Fi 5.-Semidiagrammatic dorsal view of Macrobitus x220; trunk and leg muscle not shewn, b, brain; 1g daw gland; deg, dorsal excretory gland; c, esophagus; ex, eyespot: |, leg: me, mature gg in sacihe layne. (Greatly modified from Marcus, 1 tloaca facilitate egg deposition and defeca fin. A few species have longitudinal fibers ‘sociated with the stomach, Nervous system. The lange dorsal brain fommonly has two long lateral lobes and two three median lobes; it covers much of the pharynx (Fig. 8). Two broad circum Plaryngeal connectives pass around the larynx and unite with a subpharyngeal aglion. In the trunk there is a chain of four fonspicuous ventral ganglia united by wo mt, Malpighian tubule; o, oviduct, stomach; sr, minal receptacle; sg, salivary gland; sp, sucking longitudinal nerve strands. Frequently these ganglia are slightly bilobed. Paired nerve strands originating in the brain and ganglia innervate all parts of the body. Some of the longer strands, especially those extending into the legs, have small terminal ganglionic Sense organs, The great majority of tardi- grades have a pair of eyespots in the lateral lobes of the brain. Each consists of a cup shaped mass of black or red pigment granules.: : TARDIGRADA Fic 7.~Trunk and leg musculature of Fk View. (Greatly modified from Marcus, 1998, Head cirti are usually tactile, but there is ome question about the presumed sensory junction of body cirri and filaments Cuticular structures. Fresh-water species ssually have a thicker cuticle than marine species, and semiaquatic species have a thicker aiicle than aquatic species. The surface may nooth or variously sculptured, granular orpapillate. Often there are long cirri, Some wecies of Eucardigrada have transverse ooves or folds, but Fekiniscus ane iehiniscus have dorsal armorlike thickened plates Beginning at the anterior end, these ateas follows (Fig, 12F head plate, shoulder 259 A, dorsal view; B, ventral plate (first trunk plate), a small median first intercalary, second trunk plate (divided longi tudinally), second intercalary, third trunk plate (usually divided longitudinally), third imercalary (of and the anal n_ indistinct) usually groove Many marine species are nothing short of spectacular in the development of very long cirri, lateral trunk expansions, dorsoventral Mlauening, padded daw tip: great caudal processes, and other embellishments, Reproduction, growth. The bulk of a population is always composed of females, and males reach their peak of abundance_ 260 FRESH-WATER INVERTEBRATES OF THE UNITED STATES Ventral view of nervous system of aero biotus 220. e, eyespot Il, lateral lobe of brain; Ins, longitudinal nerve strand; 3g, subpharyngeal gan sion; vg, ventral ganglion. (Modified from various during the winter or early spring. Males are unknown in Parechiniscus and Echiniscus, and these genera probably reproduce exclusively by parthenoy Some species show sexual dimorphism in the shape and cuticular struccure of the last legs, in the size and curvauure of the claws, and in the smaller size of the males. ‘The gonad is an unpaired sac dorsal to the stive tract and with one or two attachment fibers at the anterior end. In the male there are two vasa deferentia that curve around each side of the intestine and have a small swollen portion that acts as a seminal vese The female has a single oviduct on the right side of theintestine. On the other sidea bind seminal receptacle may sometimes be found in Hypsibius and Macrobictus, especially in the ate fall, winter, and early spring. In the Eutardigrada the genital duets open into the rectum, which i therefore more appropriately igrada there isa separate preanal genital pore Two distinc types of eggs have been observed in some species, thin shelled and thick shelled, and perhaps they are similarto he achyblasticand opsiblasticeggs produced by gastrotrichs, or the summer and winter a doaca. In the Heterotardi eggs produced by rotifers. At least there i some evidence to show that thick shelled eggs are produced when environmental conditions arc unfavorable; there is the further possibil that thin shelled eggs are parthenogenetic and thick shelled eggs are fertilized. The peak ofthe reproductive period is from November to May, butfemales with eggs may befoundat any time of the year In most of the true aquatic species “exter nal” fertilization is the rule. One or more males clamber about on the body of a female before she is ready to release her eggs bu after the old cuticle has become loosened in preparation for a molt. The males release sperm through the cloacal aperture or genital pore of the old cuticle, and the eggs are fertilized when they are released into the cavity beeween the new and old cuticle 1 semiaquatic species fertilization is “internal” and does not necessarily occur justbeforean eedysis. Sperm enter the genital pore or cloacal aperture and fertilize the eggs before their release from the body of the Inypi female. Sometimes the sperm may be stored in the seminal receptacle Most female Echiniscoidea produce onl to 6 eggs, but in other groups a female produces anywhere from one to 30 eggs Sometimes the whole complement may be released in 15 to 30 minutes In Macrobiotus and some species of Fypsibis the eggs are deposited freely and singly, orin groups. In other fresh-water forms they are ‘contained in the newly shed cuticle, Free eggs are often sticky for attachment to the substrate and frequently faceted, spinous, tuberculate or variously sculptured. Eggs released into the Ins ple ing ind of by: fro rep det pet dui afia TARDIGRADA the old cuticle usually have a smooth surface side the true shell a thin membrane sur rounds che developing embryo. Depending on the species and ecological conditions, a thin shelled egg usually com pletes development in $ to 12 days and a thick shelled egg in 10 to 14 days Theyoung emerge from the byruptur e shell with their stylets. Newly hatched individuals are one third to one fifth the size (mature specimens, and all growth occurs wenlargement of cells already present. Asidc fom their smallersize, immature ardigrades may be recognized by their underdeveloped reproductive system, slight differences in the deuils of the buccal apparatus, and some times by the fact that there are only two claws per leg, indicate 4 to 12 moles history, and sexual maturity is asualy attained sometime after the second or third molt. The old cuticle is usually shed ater a period of growth, but it may also be Fic. 9~Cast eae Misr igrade exoskeletons contain m (four of five eggs in p 261 shed as aresponseto hunger and unfavorable environmental conditions. Several days before jolt the sclerotized portions of the buccal apparatus are ejected and new ones (attached to the old muscles) promptly begin to formed by the salivary glands. A day or two beforeamolt, theanimal shrinks temporarily and slightly. Then the old cuticle ruptures at the anterior end and the animal crawls out hhe lining of the rectur: and the relost at molting laws are secreted by special claw glands in the tip of each leg. There appears to be considerable variation in longevity, assuming that the animal is continuously active, Two months and 24 months are perhaps the extremes, General ecolog biotus and Hypsibius ave rly aquatic and some .quat but the ecies in these genera are semi: aquatic only and occur chiefly on terrestrial A few species of Macro. are both majority of and semiaquatic WEES. A, dorsal view of Preudechiiscus; B, lateral view of cess of hatching} ‘A modified from Ramazotti and Maucci),262 FRESH-WATER INVERTEBRATE! mosses and lichens, Miluesivin tardigradum occurs in both semiaquatic and aquatic habitats, All ofthe species in the other fresh- water genera are semiaquatic and occur in true aquatic habitats only rarely and for uitously. Active tard nd if one searches diligently in masses of filamen tous algae, washings of higher aquatic plants, or in the debris on the bottom of ponds an¢ pools. The semiaquatics des can usually be fou cies, h active only when the plant has been wetted by rain or when it is splashed at streamside Mar nngiosperms contain large tardigrade pop lations, cs, liverworts, lichens, and rosette Mosses on rocks or on the bark of wees contain tardigrades, even though they may be wet enough to allow them to be active for les than 5% of the time. Concentrations up to 000 dry and inactive anabiotic tardigrades per gram of dry moss have been reported. Mosses with esp walls, such as Pubirichum, harbor tardigrades. Presumably their stylets cannot penetrate such walls, Favorable por tions of sandy beaches sometimes contain 300 to 400 tardigrades/10 cm? of wet sand Sometimes they occur as deep as S cm inthe sand. Most species appear to be eurythermal \d are normally active anywhere between wo 30°C. Some forms have been found in warm springs where they are continuously exposed temperatures of 40°C Although aquatic species are generally characteristic of the litoral, there are record of collections as deep as 100 m on lake bottoms. Semiaquatic species occur from the tropics to the Arctic and Antarctic and from scalevel to mountain tops up to 6000 m high. The annual period of activity varies corre spondingly, from the year round to only a week or two, Litle is known about minimal oxyger requirements, but it is usually inferred that tardigrades cannot endure low dissolved iny other oxygen concentrations as do ma small aquatic metazoans. Aquatic species are usually mostabundant between January and May, and sometimes there is a second maximum in the autumn OF THE UNITED STATES months. Moss dwellers donot have pro nounced seasonal maxima, Individuals ar active as long as the moss is damp. Tardigrades are chiefly preyed on by amoeboid protozoans, nematodes, and pe haps by each other to a certain extent Anhydrobiosis. When a bit of moss dries, many of the associated Tardigrada have the spusual ability o assume an inactive anabiotic ryptobiotic) state that may persist foraslong as four to seven years. When normal eco conditions again prevail come out of their quiescent state a rdigrades nd resume normal activity. Most marine species and true s apparently do nat have the ability to assume an anabiotie aquatic fresh-water spec nder drying conditions the head, posterior end, and legs are retracted, and the whole body becomes more or less rounded into a tun.” Considerable water is lost from the ody cavity and the anabiotic animal is there ore shriveled and wrinkled (Fig. 10). Interna 10.Dorsal view of a tardigrade “tun” i the anhydrobiotie state, X00, Althe for b disso able Tuns proc hum M butd toab for 6 Ri Usus hour eco high food % cally Hypsi hick the a wall A Macr have TheTARDIGRADA isceral structures are not greatly modified. Although drying ss almost always responsible jor bringing on anabiosis, a lack of sufficient dissol Tr process occurs slowly, and when the relative d oxygen and perhaps other unfavor onditions may also be responsible is can be formed only when the drying hurnidity is high Mecabolic processes proceed very slowly, butthc length of time a specimen can remain mntinuous anabiosis depends on the Death amount of stored food in the bod the supply is exhausted Anabiotic tardigrades are highly resistant iain viable even after being ke experi- mentally in strong brine solutions, at 100°C for6 hours, and at 190°C for as long as Revival from anabiosis takes but a shor time when individuals are properly wetted. wally the interval is 4 minutes to several nurs, and occasionally a day or wo. The animal absorbs water, swells, and prompdly mes active, Reviva higher temperatures, with higher dissolved xygen, and in animals having more stored fod material. Moss species usually go throug! nerous alternating periods of activity and amabiosis during their life cycle, depending nthe nature of their habitat. One author states that the life cycle may be as long as 60 Jars, including anabiotic periods Laboratory nimals can easily be dried and revived 10 Cysts. Damage, hunger, and other ab- otal ecological conditions sometimes bring about the formation of resistant cysts, espe lyin truc aquatic species of Macrobiotus and rinkled cuticle and forms a dark-colored nternal isundergoa variable degree of d «al conditions arc favorable ruptures, and the tardigrade emerges. A “simplex” stage has been observed in many tardigrades, especially in species of ribiotus. Such animals are inactive and fave amore or less reduced buccal app; Thc macroplacoids are abortive, the esopha very slender, and onal and redu 263, e siylets are no! digestive system anterior t9 the stomach disappears and there is no mouth. Litle is known about the factors responsible for the appearance of the simplex stage, but it is Jnt that it may oceut in animals about co Anoxybiosis, A third ype of inactive dition in tardigrades is called anoxybiosis. It ly comes about when there isinsufficient oxygen. The body swells and becomes rigid and turgid, There are no m0\ ements, and itis, ult to decide whether the animal is aliv. After a maximum of about 5 days in this tion the animal dies, but if food and oxygen are supplied promptly the animal indiv cr of late iduals en Sometimes ng from anabiosis go into owe (1975) recognizes two other kinds ncy among tardigrades, both n only superficially investigated. One is cryobiosis, which is closely associated with tun formation and isa state of suspended animation assumed on fre ig and other exceptionally low temperatures. The second Greatly modified fr semidiagrammatic, m Marcus, 1928,is osmobiosis, a cryptobiotic or tun state induced by exceptionally high osmotic pres sures when the animals are placed in abnormal "Everitt (1981) worked on tardigrade popu lations of an Antarctic pool that was frozen solid for 8 months each year, The animals (Hypsibius) overwintered in a “cryptobiotic State” and quickly became active in an algal mat when the pond thawed. Densities as high as 470 animals per gram of algae were found. Geographical distribution, dispersal The great majority of tardigrades ave un. doubtedly widely distributed, and an abun dance of species first found in Europe have since been found scattered over a broad range. Some are Holarctic, generally distrib uted in the Northern Hemisphere, or even cosmopolitan (Macrabiotus hilflandi Schultze, Hyp iauseri (Doyere), and Miluesm been reported only from single mainland or adum). Other species, of course, have island localities. The tropics contain notably few species and few individuals, Passive distribution is effected by currents, animals (usually insects), waves, and wind, especially in the anabiotic and cyst stages. There is considerable positive evidence for wind as an important distributive agency Collecting. Almost any bit of suitable ‘moss will yield tardigrades, Ifthe mossis wer, simply rinse it out in water and examine the washings. Masses of filamentous algae and rooted aquatics may be treated similarly. Dry mosses and lichens should be soaked for 30 minutes to several days. At intervals rinse such plant material, and the tardigrades will be found as they emerge from cysts and the anabiotic state. Culturing. Tardigrades are not easy 1 culture, and litle actual culture work seems ‘0 have been done. They may be kept in masses of filamentous algae or wet moss for a few days to weeks. Change a large fraction of the water every three or four days. Preserving, preparing. Unlike many ‘other small fresh-water metazoans, tardigrades a 264 FRESH-WATER INVERTEBRATES OF THE UNITED STATES arc casy to fix in extended form. A lang percentage will be suitable when killed and fixed in 5% formalin or 85% alcohol, Especially fine specimens may be secured by inducing anoxybiosis ‘under a cover slip sealed with petroleum jelly before killingand fixing. This procedure usually requires 10 minutes to $ hours. Some investigators recommend the use of one of a variety of narcotics before fixation. Fixatives thar are particularly useful for staining and histological work are osm acid, saturated mercuric chloride, and cetic acid. Piercing the body wall with a fine needle permits beter staining and dey dration, Simple unstained permanent mounis may be made in glycerin jelly The sclerotized parts of the buccal apparz tus may be isolated on a slide by the judicious use of 10% potassium hydroxide Taxonomy. The phylogenetic affinities tardigraces are highly debatable. During eatly ‘embryology five coelomic pouches are forme from the mesoderm, but only one poster pair persists as the gonads and their ducts the others degenerate. The main body cavity is therefore probably a hemococ. Often they are placed near the mit Phylum Arthropoda, especially be their indistinct segmentation, four pairs of legs, and piercing stylets, Some workers cow inthe sider them crustaceans, and a further sugges tion places them near the Onychophora Other workers, however, consider the Phylum Tardigrada one of the wormlike groups of various affinities and place it after the Nematoda and either before or after the 1d Bryozoa in the phylogenetic Chaetognathaa series. For the time being and until more definite evidence is forthcoming, this last suggestion is the one followed in the p ual About 50 genera are recognized in the whole phylum of which 80 contain a total o about 400 aquatic and semiaquatic fresh water species, the others being marine. Four tcen of these genera have been reported from the United States, The chief taxonomic structures used in generic and specific determinations include details of the buccal apparatus (especially macroplacoids), armour, claws, cirri, file ments, and eggs. Echiniscus, Macrobiotus, and H di de fal di th he plTARDIGRADA 265 Hypsibius are large genera that are often sub divided imo species groups or subgenera for taxonomic convenience. Most of the earlier descriptions are insufficient, owing to. the false supposition that the structures of the daws alone were sufficient and trustworthy characters, The Ramazotti and Maucci mono- graph (1983) is the most complete reference. The following key includes data for al valid fresh-water genera of the United States An even greater number of genera is known from other parts of the world. Fic. 12.-Structure of Heterotardigrada. A, dorsal view of Oreela, 280, a rare genus, not yet found in the United States; B, Par 1040; G, do of Preudechinis bead plat; il plate (divided od from various sources. ses chtonides, X430; C, claws of Echniseus,D t0F, typical species of Fk 340, Armor sculpruring omitted on D, F, and G. intercalaries 1, Firsttrunk plat; (2, second trunk plate (divide ‘nal plate hy ‘8, third trunk1 buccal structure of Hy FRESH-WATER INVERTEBRATES OF THE UNITED STATE! KEY TO GENERA OF TARDIGRADA ar separate but similar claws on Class HETEROTARDIGRADA, ‘Order ECHINISCOIDEA, ECHINISCIDAE, 2 Head without anterior cirri and lateral filaments (Figs. 140-Q); each leg with two double claws oro igs 14C-E, LN}; semiaquatieand aquatic ... Class EUTARDIGRADA, ‘Order MACROBIOTOIDEA, 3 variously sculptured; witha variable numberof cirri and filaments Figs Head with anterior cirri (rarely absent) and lateral filaments; with ‘each leg (Fig. 12): semiaquatic unlike pairs of claws Body with thick dorsal plat 12D-G); eggs deposited in shed exoskeleton (Fig. 9) 3 Body with thin dorsal plates (Fig. 12); up to 200 jum long; one species reported from Utah. arechiniscus Armor consisting of head plate, three trunk plates, and anal plate (plus intercalaries) Figs. 12D-F)iup to 830 yum long; about 100 species worldwide Echiniseus Anmor consisting of head plate, four trunk plates, and anal pate (plus intercalaries) (Fig. 12) 4 With one very long median dorsal seta; uncommon. Hypechiniscus Without such a dorsal seta; up t0 500 pm long; about 30 species worldwide. Pseudechiniscus Each leg usually with ewo heavy double claws Figs. 14C-E, LN); sucking pharynx with micro- or tnacroplacoids (Figs 3, 4); eyes present or absent . Each leg with two long, slender laws and «wo short, heavy claws (Figs. L4A, C); sucking pharynx ‘longated and without placoids (Fig. IMB); mouth surrounded by six prominent papillae up 0 1200 uma long: one cosmopolitan species... MILNESIIDAE, Milnesium tardigradum Doyére Buccal wibe with longitudinal anteroventral lamina (Fig. 13) MACROBIOTIDAE, 7 ge y +4 \\ H i\} a |] c De Fats a rT) . yD tte al serucnure and claws of tardigrades. A, buccal structure and claws of Macrbotay ss C, buceal structure of Preudediphascon; D, claws of Dachebius: E, claws of , buccal surueture of Dipkascn, H, buceal structure fron 1m Schuster et al, 1980. Fic, 18.-Bug pbs F claws of Fy uct hristenberry and Higgins, 1979; all others fr Bi ‘anti cee Mocxg of Fy ‘ARDIGRADA 267 Sa —N Fic. 14Structure of Macrobiotoidea. A, Mi gradu: C, claws of M,tardgradum,; D and E, claws of Me bus: Lwo N, claws of fypribius O to Q, an, X75; B, buccal apparatus of M, us Fand G Hk, sal view of typical species of Hypribis, X20, Buccal tube wit such a lamina (Fig. 138) HYPSIBIIDAE, 11 Claws single, although occasionally a basal minute spur is prese also; 2 single species known only from Sonora, Mexico, and CA Haplomacrobiotus hermosillensis May Each claw normal and double, either similar or dissimilar in size 5 Buccal tube long and thin, with spiral thickenings Fig. IC); reporved only froun Al Pseudodiphascon Buccal tube of the usual constructc 7 Fourth legs rudimentary; in wet mosses, one species reporied from GA exapodibius Fourth legs normal in size; eggs mostly free and sculptured (Figs, 148, G) 10 Both branches ofeach claw with che tips close together (Figs. 14D, E); up to 1100 pamlongs about 100 species Macrabiotus Tips of two claws of each leg remote (Fig. 13); uncommon Dacrylobiorus Claws of similar shape and size; branches solidly or broadly joined (Fig. 138), about 80 worldwide cies but seldom reported from the United States, Isohypsibius Claws dissimilar in size and shape, one branch with thin articulation (Fig. 13F 2 uccal tube short, rigid (Fig. 188); about 30 species worldwide Hypsibiu Buccal tube long, flexible, with spiral thickenings (Figs. 136, H 1s Pharynx with normal placoids; 45 species worldwide Diphaseon Pharyngeal placoids reduced or absent Iraquascon CCFRESH-WATER INVERTEBRATES OF THE UNITED STATES. 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