Chapter II: Part 2
Body spherical to pyriform, constricted near the middle, the constriction dividing the body into dissimilar parts. The anterior part is broadly pyriform, somewhat plastic and hyaline, with an oral extremity which is sometimes hollow, sometimes evaginated and convex. Upon this flexible anterior part there are four short but distensible tentacles. The posterior part is granular and usually filled with food particles; it is well rounded and holds the nucleus and contractile vacuole. The entire body is surrounded by a fine cuticle. The nucleus is elongate and extends through the greater part of the posterior half. The contractile vacuole lies on one side, near the girdle. The mouth is on the anterior pole in the tentacle region. The motile organs are cirri and cilia, all inserted in the constriction. There are two sets of cirri and one of cilia; the latter stand out radially from the girdle and are usually in motion. The cirri of one set, the anterior, extend forward about twice the length of the anterior half; those of the posterior set closely engirdle the lower half, reaching not quite to the posterior extremity. These are somewhat hyaline and are closely approximated, giving the impression of a tight-fitting crenulate casing about the lower half. The cirri are sharply pointed, much broader at the base, and the two sets are so placed that, looked at from above, they have the appearance of a twisted cord. (Fig. 31 b.) Movement erratic; sometimes the animal swims steadily forward with mouth in front; again it shoots across the field of the microscope, either backward or forward or sideways, through the action of its powerful cirri. It is often quiet, usually mouth downward, and is held in place by adhesion of the tentacles. In this position it looks strikingly like a heliozoön.
Length 35µ; greatest width 30µ. Not uncommon.
The chief features by which this species is distinguished from the frequently described _M. pulex_ of Europe are the number of anterior cirri and the ring of true cilia in place of the central girdle of cirri. The European form is described with four anterior bristles; the present form has from 28 to 32. The radial cilia differ decidedly from the more powerful cirri and they are not in one plane, so that counting is difficult; they are not closely set. The presence of tentacles makes these forms of especial theoretical interest, especially in the light of the origin of _Suctoria_.
Genus TIARINA R. S. Bergh '79.
(Claparède & Lachmann '58.)
Body subcylindrical, pointed posteriorly, two and one-half times as long as broad; encased in covering composed of separate pieces arranged in five girdles. The pieces bear processes which rest against neighboring pieces of the girdle. Mouth large, anus terminal near contractile vacuole. The macronucleus is simple and round. Salt water.
Tiarina fusus (Cl. & Lach.) emend R. S. Bergh.
Synonyms: _Coleps fusus_ Cl. & Lach. '58; Daday '86; Möbius '88, Lauterborn '94; Shevyakov '86.
This form, which resembles _Coleps_ rather closely, was placed as a separate genus by R. S. Bergh. The skeletal parts consist of five zones of needles composed of an organized substance and embedded in the cortical plasm, the last zone coming to a point at the posterior end. The needles have lateral processes, which give a latticed appearance to the casing. The cilia are long, with a specialized crown of still longer ones at the oral end; they arise outside of the skeletal elements and do not pass between them, as in _Coleps_.
KEY TO MARINE GENERA OF TRACHELINIDÆ.
Diagnostic characters: Body bilateral, or asymmetrical by local prolongations; usually compressed or flattened laterally, the left side more convex than the right. The essential feature is the position and character of the mouth. This is either a long slit extending from the anterior end well down the ventral surface, or the posterior part only of a ventral furrow remains open as a round or elongate mouth some distance from the anterior end. The entire mouth region of the body is usually drawn out into an elongate tapering proboscis which is generally curved dorsally at the extremity. An oesophagus is short or absent altogether; when present it is supported by a stiff buccal armature. Cilia are uniform about entire body or limited to the flat right side. Food is swallowed.
1. a. Proboscis easily distinguished 2
from the main body
b. Proboscis not marked off from Genus *_Loxophyllum_
main body; body flat;
both surfaces striated
2. a. Mouth runs the entire length Genus _Amphileptus_
of proboscis; entire body
uniformly ciliated
b. Mouth runs the entire length Genus *_Lionotus_
of proboscis; body flat; right
side only is ciliated
c. Proboscis much drawn out, Genus _Dileptus_
flexible; mouth at its base
* Presence at Woods Hole indicated by asterisk.
Genus LOXOPHYLLUM Dujardin '41.
(Duj. '41; Wrzesniowski '69; Quennerstedt '65; '67; Cohn '66; Entz '84; Gourret & Roeser '88; Bütschli '88; Shevyakov '96.)
The body is flat and somewhat leaf-shape, flexible, and elastic. The anterior end is somewhat proboscis-like and flexible, but is not sharply demarcated as in _Lionotus_. The central portion of the body is developed into a more or less arched dorsal mass, which usually contains the nuclei and contractile vacuoles. As a result of this local thickening, the body is surrounded by a thin hyaline margin. This, however, may be absent on the right side in some species. The mouth reaches from the anterior extremity to a short distance from the end, and usually approaches the left edge. An anus is present near the posterior end of the dorsal swelling. Trichocysts are numerous on the ventral surface, and often on the dorsal surface, where they are inclosed in minute papilla-like swellings. Cilia-distribution controverted. Maupas and Bütschli hold that ventral surface alone is ciliated; others (Kent and Dujardin) that cilia are uniformly distributed. The entire body, dorsal and ventral surfaces alike, are uniformly striated. The contractile vacuole lies posteriorly, on the right side and in the dorsal swelling. In the fresh-water form _L. meleagris_, it is connected with a long canal whose swellings are frequently taken for additional contractile vesicles (Bütschli); in the marine form described below the canal is not developed and a series of vacuoles takes its place; these are all contractile. The macronucleus may be single, double, quadruple, band-formed, or rosette-formed. Movement is steadily progressive and peculiarly gliding. Fresh and salt water.
Loxophyllum setigerum Quenn. '67.
Synonyms: _Litosolenus armatus_ Stokes '93; _Litosolenus verrucosa_ Stokes '93.
The body is flattened, irregular in outline, obtusely pointed anteriorly, the point being turned to the right; rounded posteriorly. The left edge is nearly straight, the right considerably arched with a few setæ on the posterior half. Contractile vacuoles are numerous, dorsal in position and on the right side. The macronueleus is beaded, the several spheres connected.
Variety armatum (Cl. & Lach.) Fig. 32.
Under the name _Litosolenus armatus_, Stokes described a form from brackish water near New York, which should unquestionably be referred to the genus _Loxophyllum_, and I believe to Quennerstedt's species _setigerum_. While the latter possesses only a few setæ, the former has a number of them, and Stokes described his species as having a variable number. For this reason I include the Woods Hole form under the tentative name _armatum_, as a variety of Quennerstedt's _L. setigerum_. The flat margins are distinctly striated longitudinally, and faintly marked radially, on the dorsal surface. Longitudinal elevated striæ also run the length of the dorsal hump and upon the entire ventral surface. The ventral surface is alone ciliated. Upon the edges of the flat border are sharp-pointed, colorless, spine-like processes, situated at equal distances around the entire periphery except at the anterior end. Each spine is thick at the base and tapers to a full point which is curved upward--_i. e._, dorsally (fig. 32, a, b). The entire body is plastic and contractile, turning its leaf-like edge readily over objects upon which it creeps. The cilia are fine and uniform, with a tendency to lengthen in the oral region.
Length 100µ; greatest width assumed on contraction 85µ; when normal about 50µ.
Genus LIONOTUS Wrzesniowski '70.
(Incorrectly called _Litonotus_ by many. Entz '84; Gruber '84; Bütschli '88; Kent '81; Schewiakoff '89; Shevyakov '96.)
The body is elongate and somewhat lance-shaped, widest at the central part and tapering to a point at the anterior end. The posterior end may be similarly tapered or rounded. The anterior end frequently proboscis-like, flat, and flexible, while the entire body is more or less elastic and contractile. The right side is flattened and alone provided with cilia, while the left side of the body proper is arched; on the left side of the proboscis is a row of coarse cilia resembling an adoral zone, and a row of trichocysts. A long peristome stretches down the thin, ventral side of the proboscis, and the mouth proper is situated at the junction of the proboscis and body; the mouth, as a rule, is invisible. The ciliated right side alone is striated in the majority of species. The contractile vacuole may be single or multiple, usually in the posterior region of the body and dorsal in position. The macronucleus is usually double, rarely single or quadruple, but may occasionally break into numerous smaller pieces. Movement, free-swimming or gliding, with especial tendency to get under clumps of foreign matter.
Fresh and salt water.
Lionotus fasciola Ehr. Fig. 33.
Synonyms. _Amphileptus fasciola_ Ehr. '38; Dujardin '41; Lachmann
'56; Cohn '66, Diesing '65.
_Loxophyllum fasciola_ Claparède & Lachmann '58;
Balbiani '61.
_Loxophyllum duplostriatum_ Maupas '83. Shevyakov '96.
Body frequently brown or brilliant yellow in color, somewhat sigmoid in form with tapering anterior end, the extremity of which is turned dorsally. The proboscis is about half the entire length and is not sharply marked from the rest of the body but tapers gradually, its base being equal to the diameter of the body at its middle point. The body is slightly contractile and the posterior end is carried to a rounded point, but not into a distinct tail. Unlike the fresh-water variety, this one has no hyaline margin nor hyaline caudal region, and the contractile vacuole is double or multiple on the dorsal side near the posterior end. Cilia are present only on the under (right) side, with, however, a row of large cilia marking the course of the elongate mouth, upon its left side. The right side is striated, the left arched and without markings. The endoplasm is finely granular with, however, larger food particles in the process of digestion, while specimens are occasionally seen with the natural form completely lost through distortion caused by over-large captures (Cf. also Wrzesniowski '70, p. XXIII, fig. 32). Movement continuous, slow, and gliding; very little tendency to jerking movements. Macronucleus double, both parts spherical, and placed in about the center of the larger part of the body; closely approximated but not, as Schewiakoff described, connected. In conjugation, a large form unites with a smaller one, the mouth parts being connected. Details of conjugation and macronuclei not made out. Length 200µ to 600µ.
KEY TO MARINE GENERA OF CHLAMYDODONTIDÆ.
Diagnostic characters: Form usually ellipsoid, never very elongate. Transverse section of body circular or elliptical. The mouth is usually some distance from the anterior end and may be in the posterior part. Sometimes it is in the center of the ventral surface, again on the right side. The oesophagus invariably has a well-developed buccal armature, or a smooth peculiarly built oesophageal tube. Food particles of large size.
1. Body cylindrical. Cilia about Genus *_Nassula_
entire body
Body flat 2
2. a. Without a caudal process 3
b. With a caudal process 5
3. a. Anterior end angular 4
on left side
b. Anterior end rounded Genus _Chlamydodon_
4. a. Dorsal striæ and cilia present, Genus _Orthodon_
ventral cilia longer
b. Dorsal striæ and cilia absent; Genus *_Chilodon_
posterior end not pointed
c. Dorsal striæ and cilia absent; Genus _Scaphidiodon_
posterior end pointed
5. a. Caudal spine with posterior 6
bristle-like cilia
b. Caudal spine without posterior Genus _Trochilia_
bristle-like cilia;
ventral cilia reduced
6. a. With pigment spot on anterior Genus _Ægyria_
angle
b. Without such pigment spot Genus _Onychodactylus_
c. Cilia on right edge only of Genus *_Dysteria_
greatly reduced ventral surface
* Presence at Woods Hole indicated by asterisk.
Genus NASSULA Ehr. '33
(Dujardin '41; Stein '67; Cienkowsky '55; Cohn '66; Clap. et Lach. '58; Kent '81; Maupas '83; Entz '84; Fabre-Domergue '88; Bütschli '88; Shevyakov '96.)
The body is ovoid or cylindrical, with well-rounded ends, and in some cases slightly flattened. The mouth is ventral and placed some distance from anterior end (1/4 to 1/3 total length). A slight depression on the ventral surface marks the mouth region, which is further indicated by larger and more powerful cilia. The rest of the body is uniformly ciliated. The entire body is marked by clearly defined spiral stripes. The mouth is circular and the oesophagus is supported by a considerable armature, which usually extends dorsally and to the left, rarely to the right. In some cases the structure of this armature is indistinct; again it can be clearly seen to consist of definite rods (Stäbchen). The anus is probably always terminal. Contractile vacuoles are variable in different species. In some cases there is but one, which is placed at the posterior end or centrally on the ventral side; in others there may be four--two dorsal and two ventral. In many cases trichocysts are uniformly distributed. Sometimes the body is colorless; again, and more often, it is brightly colored with red, blue, brown, or black pigment. The macronucleus is globular and central, occasionally band-form and with numerous attached micronuclei. Food substance varied, usually vegetable matter, see, however, below. Cysts are globular. Movement is a steady progression, combined with rolling.
Nassula microstoma Cohn '66. Fig. 34.
Synonyms: _Paramoecium microstomum_ Cl. et Lach. '58, Gourret et Roeser '88; _Isotricha microstomum_ Kent '81.
Body subcylindrical, rounded at each extremity, not quite twice as long as broad. A slight depression on one surface marks the position of the mouth, this depression being indicated by a row of longer cilia. The mouth is extremely small and is surrounded by a curious buccal armature. This is not made up of bars or rods, as in most species of _Nassula_, but appears perfectly smooth and uniform except for the considerable swelling at the inner end. The cuticle is firm and unyielding and marked by longitudinal and somewhat spiral rows of cilia and trichocysts. Under the microscope this is one of the most pleasing forms found at Woods Hole. Its color is yellowish brown from the presence of brilliant particles of coloring matter held in the cortical plasm, and, as it slowly rolls along, these particles and the black trichocysts give to the organism a peculiar sparkling effect. The macronucleus is almost central; the contractile vacuole posterior. The endoplasm appears well filled with food bodies, some of which could be distinguished as _Amphidinium_ and _Glenodinium_.
Length 55µ; greatest diameter 30µ.
Genus CHILODON Ehr.
(Dujardin '41; Engelmann '78; Stein '54, '58; Kent '81; Bütschli '88; Gruber '83; Cienkowsky '55; Möbius '88; Clap. et Lach '58; Wrzesniowksi '65; Shevyakov '96.)
Small forms, greatly flattened dorso-ventrally and almost egg-form in outline. The anterior end is bent distinctly to the left and forms a characteristic process, which, together with the entire margin of the body, is soft and flexible. The posterior end is, as a rule, broadly rounded. The ventral surface is finely striate, and this surface alone is ciliated. The lines of cilia converge at the mouth, and at this region the cilia are somewhat larger and more distinct, thus forming a functional adoral zone. The mouth is median and is situated in the anterior half of the body. It is surrounded by a well-defined armature, composed usually of from 10 to 16 rods. The contractile vacuoles are quite varied and from one to many in number, the number increasing with the size of the individual. The macronucleus is usually single, elliptical in form, and centrally placed; one micronucleus. Reddish granular pigment and trichocysts are occasionally present.
Chilodon cucullulus Müll., sp. Fig. 35.
Synonyms; _Colpoda cucullus_ O. F. Müller; _Loxodes cucullulus_; _Chilodon uncinatus_ Ehr. '58, Perty '52, Dujardin '41; _L. dentatus_ Duj., etc.
This extremely variable form has received so many different names that it hardly pays to enumerate them. It is one of the commonest and most widely spread ciliates known, although at Woods Hole I was surprised to see it so rarely. It is the type species of the genus and needs no further description. The specimens observed at Woods Hole had numerous contractile vacuoles and were 42 to 45µ long and from 28 to 32µ wide.
Genus DYSTERIA Huxley '57.
(Cl. et Lach. '58; Entz '84; Möbius '88; Shevyakov '96.)
Small forms, firm in outline, and colorless or slightly colored. The body is somewhat clam-shaped, flattened, slightly curved or straight on the right side, the other more convex. The true ventral side is only a narrow strip along the right and anterior edge of the body, the apparent ventral side being a fold of the very large dorsal surface which comes around ventrally, forming a valved structure somewhat analogous to a clam shell. Cilia are limited to the outer edge of the small ventral surface, which also bears a peculiar spine at the posterior end. Behind this spine are larger cilia. The mouth opening lies in the anterior widened portion of the ventral surface and is connected with a smooth tubular pharynx. The right half of the dorsal side, _i.e._, the apparent dorsal side, is arched and bears longitudinal ridges. Two to four contractile vacuoles are placed on the ventral side. The macronucleus is usually dorsal, elliptical, and cleft, with one micronucleus attached. Fresh and salt water.
Dysteria lanceolata Cl. et Lach. Fig. 36.
Synonym: _Cypridium lanceolatum_ Kent '81.
Outline of the flattened body ovoid; body consists of two valve-like portions; the edge of the right valve is nearly straight, that of the left valve more or less sinuous; anteriorly it is cut away, obliquely and posteriorly it has a deep indentation in which the seizing spine rests. The cilia are confined to the ventral surface, here reaching, however, from the anterior dorsal extremity to below the posterior indentation. Posteriorly the cilia become larger, corresponding to the larger cirri of _D. armata_, which are posterior to the spine. The mouth lies between the two valves and is surrounded by a long and smooth buccal armature which passes downward and backward to the left a distance equal to about half the entire body length. The macronucleus is situated in the dorsal region in the central part of the body. There are two contractile vacuoles, one behind the center of the buccal armature, the other near the inner end of this organ. Movement is in circles, the animal moving around quite rapidly when not attached by its posterior process. It is colorless and measures 45µ in length by 27µ in width. Claparède & Lachmann and Shevyakov describe it as 70µ long.
KEY TO MARINE GENERA OF CHILIFERIDÆ.
Diagnostic characters: Mouth never lies behind the middle of the body; the oesophagus is but slightly developed. The undulating membranes are placed either on the edge of the mouth or in the oesophagus. A peristomial depression leading to the mouth is absent or very slightly indicated.
1. Mouth in the anterior half, Genus *_Frontonia_
undulating membrane on left edge
only; right edge continued in a
long ventral furrow
2. Two undulating membranes; mouth Genus *_Colpidium_
central; no caudal bristles
3. Two undulating membranes; caudal Genus *_Uronema_
bristle
Genus FRONTONIA Ehr. (Cl. & Lach. '58?).
(Ehrenberg, subgenus of _Bursaria_ '38; Claparède & Lachmann '58; Bütschli '88; Shevyakov '96.)
Form elongate and cylindrical, or often flattened dorso-ventrally, with round or pointed ends. It is usually plastic and contractile. Cilia are evenly distributed about the body and are similar in length. The large, open mouth lies on the anterior half of the ventral surface, and is elongate and oval in outline. On its left edge is a well-defined membrane which stretches across to the right side of the mouth. On the right edge is a small, longitudinally striped tract which is free from trichocysts and smooth in appearance. This tract is continued posteriorly in a long furrow, which in some cases reaches the posterior end of the animal. A few rows of cilia in this furrow vibrate differently from the others and give the effect of a membrane (Bütschli). The oesophagus is extremely short and hard to make out. The body is usually covered uniformly with trichocysts, often of considerable size. There are 1 or 2 vacuoles with long canals radiating throughout the endoplasm. The macronucleus is oval and centrally placed. Micronuclei vary from one to many. An anal opening is placed at the end of the long ventral furrow. The plasm is colorless or green by the presence of Zoochlorella, or colored brown or black by pigments. In these cases there is a considerable pigment mass on the anterior end. Movement is regular, forward, and combined with rotation. Food consists of foreign objects, diatoms, other protozoa and the like. Fresh and salt water.
Frontonia leucas Ehr. Fig. 37.
Synonyms: _Frontonia vernalis_ Ehr. '38; _Bursaria leucas_ Allman '55, Carter '56; _Panophrys leucas_ Duj. '41, Stein '67; _Panophrys vernalis_ Dujardin '41, Stein '67; _P. chrysalis_ Duj. '41, Fromentel '74; _Cyrtostomum leucas_ Stein '67, Kent '81.
Form ovoid, elongate, occasionally a little flattened dorso-ventrally. Mouth in the anterior third of the body. The left edge of the mouth carries a distinct undulating membrane; the right edge is plain, longitudinally striated and bears cilia. It is slightly depressed and the depression is carried posteriorly in the form of a shallow furrow which reaches to the posterior end. The contractile vacuole is on the left side, the spheroidal nucleus on the right side of the furrow. The body is uniformly covered with fine cilia, and the periphery is uniformly studded with large trichocysts, except along the furrow. Food consists of dinoflagellates and other small forms. Color dark brown to black.
Length 330µ; width 200µ.
This form differs considerably from the fresh-water _Frontonia leucas_ as described by Schewiakoff '89, especially in the extreme length of the peristomial furrow, in the position of the nucleus and contractile vacuole, and in the nature of the water canals. These in the Woods Hole form are very irregular in size and very much branched, not uniform as in Lieberkühn's (see Bütschli) figure of _Frontonia leucas_, nor radiating as in Schewiakoff's description. This may be the same species as _Frontonia marina_, of Fabre-Domergue '91, whose description and figure I have not seen.
Genus COLPIDIUM Stein '60
(Bütschli '88; Maupas '83.)
The general form is oval, slightly compressed laterally with the dorsal side strongly arched. The ventral side is slightly incurved. The anterior end is somewhat smaller than the posterior end, which is broadly rounded. The mouth is placed some distance from the anterior end in an oral depression and opens into a tubular oesophagus. There are usually two undulating membranes which do not extend beyond the mouth borders. The right undulating membrane extends down into the oesophagus and appears to be attached to the walls of the latter. The body stripes in front of the mouth are twisted to the left. The anus is terminal and the contractile vacuole may be terminal or situated forwards in the dorsal region. The macronucleus is spherical and has one micronucleus attached. Food consists mainly of bacteria. Movement rapid, but interrupted.
Fresh and salt water, common in infusions.
Colpidium colpoda Ehr., sp. Fig. 38.
Synonyms: _Colpidium cucullus_ Kent '81; _C. striatus_ Stokes '85; _Kolpoda cucullus_ Duj. '41; _Paramoecium colpoda_ Ehr. '38, Quennerstedt '67; _Plagyiopyla nasula_ Kent '81, G. & R. '86; _Glaucoma pyriformis_ G. & R. '86; _Tillina campyla_ Stokes '85, '88.
The body is oval, somewhat larger posteriorly, and a little compressed dorso-ventrally. The anterior end is twisted a little from the right to the left (more evident in fresh-water forms), and leans somewhat toward the ventral side. Under this portion, on the ventral side, lies the mouth in a large depression just above the middle of the body. The entire body is covered with uniform and delicate cilia, which are placed in longitudinal rows. These rows are almost straight on the dorsal side, but bend on the ventral surface, following the contour of the twisted anterior portion. The endoplasm is finely granular; the oesophagus leading into it is very distinct. Schewiakoff ('89) describes two membranes, an inner and an outer; Maupas ('83) describes them as right and left. In the present species I was able to make out only one. The macronucleus is central, spherical in form, and bears a single minute micronucleus. The contractile vacuole is posterior and dorsal to the long axis of the body. The anus is ventral to this axis and also posterior. Length 45µ, width 20µ. Common.
This marine variety is much smaller than the fresh-water form and the form differs in a number of respects, viz, in the anterior torsion and in the structure of the mouth. These may be, however, only individual variations of a widely spread species, and I believe it is perfectly safe to describe this as _Colpidium colpoda_.
Genus URONEMA Duj. '41.
(Quennerstedt '69; Cohn '66; Kent '81; Bütschli '81; Schewiakoff '89; Shevyakov '96.)
Minute forms; colorless and constant in body form. The form is oval, slightly compressed on the ventral side, while the dorsal side is distinctly arched. The membrane is distinctly marked by rather widely separated striæ. These occasionally have a spiral course about the body; in all cases they can be easily counted. The mouth is large and placed near the center of the ventral surface. It is sometimes approached by a very shallow depression or peristome from the anterior end, and marked by two rows of cilia. An undulating membrane extends down the mouth. Oesophagus absent. A long, stiff bristle extends outwards from the posterior end. The contractile vacuole is terminal or subterminal and near the anal opening. The macronucleus is spherical, centrally placed, and with one micronucleus closely applied. Movement is rapid and usual forwards in a straight line, often found resting, however, with outstretched cilia in contact with some foreign body. Food mainly bacteria. Fresh and saltwater, usually in decomposing vegetable substances.
Uronema marina Duj. Fig. 39.
Synonyms: _Enchelys triquetra_ Dujardin; _E. corrugata_ Duj.; _Cryptochilum griseolum_ Maupas '83; _Philaster digitiformis_ Fabre-Domergue '85.
Small animals with ellipsoidal form and about twice as long as broad. The mouth lies in the upper half of the body and bears a well-developed undulating membrane upon its left edge. The membrane is longitudinally striped and covered with long and vibratile cilia. The right edge of the mouth bears cilia which are about the same in size as the body cilia, but are more closely inserted (Schewiakoff). The most characteristic feature is the long caudal bristle, which is extremely delicate and about two-thirds the length of the body. Schewiakoff thinks this bristle has a sensory function. I could not make this out, for although other protozoa ran against this bristle, often bending it well over to one side, the animal showed no sign of irritability but lay quiescent. A spherical macronucleus with attached micronucleus lies in the center of the body. The contractile vacuole is posterior in front of the bristle. The macronucleus was found to be double, as though just divided, in a large percentage of cases. This may be a precocious division of the nucleus long before signs of the body division are evident. Such a phenomenon, however, is rare, the macronucleus usually dividing at a late stage of cell division. Length 30 to 50µ; width 15 to 20µ. Common in decomposing algæ.
KEY TO MARINE GENERA OF PLEURONEMIDÆ.
Diagnostic characters: The mouth is at the end of a long peristome running along the ventral side; the body is dorso-ventrally or laterally compressed. The entire left edge of the peristome is provided with an undulating membrane which occasionally runs around the posterior end of the peristome to form a "pocket" leading to the mouth. The right edge of the peristome is provided with a less-developed membrane. There may or may not be a well-developed pharynx.
Body small; not produced into Genus *_Pleuronema_ neck-like elongation
Body medium-sized; anterior end Genus *_Lembus_ produced into neck-like elongation
* Presence at Woods Hole indicated by asterisk.
Genus PLEURONEMA Dujardin '41.
(Perty '52; Clap. & Lach. '58; Stein '59, vol. I; Quennerstedt '67; Kent '81; Bütschli '88; Schewiakoff '89; Shevyakov '96.)
Small to medium-sized ciliates, with an unchanging form. They are somewhat lens-shape and laterally compressed, the two surfaces about equally arched. The ventral surface is nearly straight or but slightly arched; the dorsal is quite convex. The anterior and posterior extremities are equally rounded. The peristome begins as a small depression, but becomes larger until it takes in nearly all of the ventral surface. The depression becomes much deeper at about the center of the body, and is especially marked on the left side of the peristome. In this deeper portion is the mouth, with an almost imperceptible oesophagus. Upon the left edge of the peristome is a high, undulating membrane, sail-like in appearance when extended. This may stretch around the posterior edge of the peristome and upon the right aide, thus forming a pocket by means of which the food particles are directed into the mouth. The rest of the right edge of the peristome is occupied by closely approximated powerful cilia (Bütschli) or a second undulating membrane (Stein). The body cilia are relatively long. Trichocysts and caudal bristles may be present. The contractile vacuole is subterminal and dorsal; it is questionable whether there are canals leading to it. A round macronucleus with one micronucleus is in the anterior half of the body. The anus is terminal. Food is chiefly bacteria. Movement combines springing with swimming and rotation. Fresh and salt water.
Pleuronema chrysalis Ehr., sp. Fig. 40.
Synonyms: _Pleuronema crassa_ Dujardin '41; _P. marina_ Duj. '41; Fabre-Domergue '85; _P. coronata_ Kent '81; _Paramoecium chrysalis_ Ehr. '38; _Lembadion orale_ G. & R. '88; _Histiobalantium agile_ Stokes '85, '88.
The body is ovoid, slightly flattened, rounded at both ends, the anterior end sharper than the posterior. The ventral surface is almost entirely taken up by a peristome which extends from the anterior end posteriorly three-quarters of the body length. The posterior end of the peristome is straight, the left curved, following the depressed portion. The body is covered with fine cilia in longitudinal lines, except on the peristome. The mouth is small and situated in the hollow of the peristome near the left border. On the left peristome edge is a large undulating membrane. It begins near the anterior end of the body and increases in height posteriorly following the peristome edge around on the right side. This posterior bend of the membrane causes the appearance of a full sail, so often seen. It can be entirely withdrawn and folded together in the peristome. On the right edge of the peristome are large, powerful cilia. The contractile vacuole is central and dorsal; the macronucleus is in the anterior half of the body, with one attached micronucleus. Food consists of bacteria. Not very common. Fresh and salt water. It often remains quiet, with membrane and cilia outstretched, as though dead, but suddenly gives a spring and is gone.
Pleuronema setigera, n. sp. Fig. 41.
Body colorless, elongate, and with the general form of a cucumber, the posterior end being somewhat pointed. The mouth and relatively small peristome are situated in the lower third of the body. The peristome begins as a shallow furrow at the center of the ventral surface and dips sharply into the buccal depression, which is deep and turned toward the posterior end. The left edge of the peristome bears a high undulating membrane, which extends anteriorly only as far as the center of the body; posteriorly it passes around to the right edge of the peristome, thus forming the characteristic membranous pocket. Inside the oral depression is a second undulating membrane, running down to the mouth. This is small and without an oesophagus. The body is clothed with long setose cilia which are frequently fully outstretched when the animal is resting, a slight tremor of the large membrane alone indicating vitality. Posteriorly these appendages are drawn out into long filiform setæ, the number varying in different individuals from three to nine or ten. These are extremely fine and difficult to see without a high power (_e.g._ 1/12 oil) and careful focussing of the substage condensor. Like _P. chrysalis_, the resting periods are terminated by sudden springs, otherwise the movements are steady and forward. The macronucleus is central, and the contractile vacuole posterior and terminal. Length 45µ to 50µ; greatest diameter 17µ. In decaying algæ.
It was this form, I believe, that Peck '95 described as a "ciliate."
Genus LEMBUS Cohn '66.
(Cohn '66; Quennerstedt '69; Kent '81; Fabre-Domergue '85; Gourret & Roeser '88; Bütschli '88; Shevyakov '96.)
Free-swimming animals of elongate form, more or less elastic, and flexible, bending readily to avoid obstacles, etc. The anterior half is usually drawn out into a slightly curved neck-like portion. The peristome is a small groove leading from the anterior end to the mouth about midway down the ventral side of the body. Bütschli, following Quennerstedt, describes an undulating membrane on each side of the peristome groove. Other observers, however, usually describe but one, the left, which is clearly defined and stretches out some distance from the body, while the right border is described as having smaller but very active cilia. The general body surface is clothed with fine, uniform cilia, and body striæ are usually absent. One or more caudal bristles may be present. The contractile vacuole is posterior and terminal, and may be multiple. The macronucleus is spherical and perhaps double (Kent). Food is chiefly bacteria, and the animals are frequently found with the anterior end embedded in zoogloea masses. Salt water, usually in infusions.
Lembus infusionum, n. sp. Fig. 42.
The body is elongate, lancet-shaped, with a tapering anterior extremity. The dorsal outline is concave through the bending of the anterior end, while the ventral outline presents an even, convex curve. The mouth lies slightly above the center of the body and marks the posterior limit of the ventral peristomial groove, which curves slightly from the anterior extremity. Each side of this groove bears an undulating membrane, the left being much larger and conspicuously striated. The general form of this left membrane is triangular, the widest part is anterior, the narrowest at the mouth. The right membrane is similar in form, but smaller and more active. The endoplasm is colorless and finely granular, not regionally differentiated. The ectoplasm consists of a relatively thick cortical plasm specially noticeable in the posterior half of the body and a delicate cuticle which bears almost imperceptible longitudinal markings--the insertion points of the fine cilia. The body is covered with uniform cilia except at the anterior extremity. Here they are much larger and bristle-like. I was unable to find any cilia in the peristome. One long caudal bristle, one-quarter of the length of the body, trails out behind. The macronucleus is spheroidal and placed near the center of the body; a conspicuous micronucleus lies near it. A row of contractile vacuoles extends from the posterior end. I have seen as many as six of nearly equal size and one or two smaller ones. The intervals of contraction are quite long. Length 70 to 75µ; greatest diameter 10 to 12µ.
_L. infusionum_ resembles _L. elongatus_ in its general form and in its mode of life, for it excavates a retreat in zoogloea masses and lies there for considerable periods perfectly quiet. It differs from _L. elongatus_ and from _L. velifer_ (probably the same as _L. elongatus_ of Claparède & Lachmann) in the presence of the caudal bristle, in the absence of annular markings, number of contractile vacuoles, and in the slightly smaller size. It resembles _Lembus verminus_ (Müller) as described by Kent (_Proboscella vermina_), and _L. intermedius_ as described by Gourret & Roeser (_Lembus verminus_ syn.)in the absence of annular markings and in the presence of a caudal bristle. It differs from the former, however, in the absence of a tentacle-like process, and from both in the absence of a double nucleus and in the presence of many vacuoles. These features are so characteristic of all the specimens examined that I have concluded, somewhat reluctantly, to give it a specific name. It is common in old infusions of algæ, especially after decomposition is well advanced. Its food consists of bacteria.
Lembus pusillus Quennerstedt 1869. Fig. 43.
Synonym: _L. subulatus_ Kent 81.
This species is much smaller than the preceding, and might easily be mistaken for _Uronema marina_. It is subcylindrical in form, the anterior end bluntly pointed, the posterior end rounded. The oral apparatus is quite different from _Uronema_. The mouth, as in the preceding species, is at the end of a long peristomial groove extending from the anterior end to the middle of the body. The edges of the peristome bear undulating membranes as in _L. infusionum_. Like the latter, there is one caudal bristle, but unlike it there is only one posterior contractile vacuole, while the endoplasm is filled with large granules or food balls. The cuticle is distinctly striated with longitudinal markings, and the cilia are uniform in length.
Habitat similar to that of _L. infusionum_, in zoogloea masses. Length 26 to 30µ; diameter 7 to 8µ.
Although Quennerstedt's description of _L. pusillus_ makes no mention of a caudal bristle, the size and other characters are so closely similar that I hesitate to make a new species. The bristle is extremely delicate, scarcely thicker than a cilium, and easily overlooked, yet with proper focussing of the condenser I found it on every specimen examined.
KEY TO MARINE GENERA OF OPALINIDÆ.
Diagnostic characters: The form is oval, and the body may be short or drawn out to resemble a worm. They are characterized mainly by the absence of mouth and pharynx.
Anterior end not pointed; body Genus *_Anoplophrya_ cylindrical; tapering
Anterior end pointed; body elongate; Genus _Opalinopsis_ cylindrical; tapering
* Presence at Woods Hole indicated by asterisk.
Genus ANOPLOPHRYA Stein '60.
(Stein '60; Claparède '60; Leidy '77; Vejdovsky '79; Kent '81; Balbiani '85; Bütschli '88; Shevyakov '96.)
The general form is elongate, cylindrical or slightly flattened, with rounded ends, the posterior end tapering. The body is striated with clearly defined, often depressed lines, which run longitudinally and sometimes spirally. The contractile vacuoles are usually placed in rows upon the edges. The macronucleus is almost always long and band-formed, rarely oval, and generally extending through the entire length of the body. Micronuclei have been made out in one case. Reproduction is effected by simple cross division or by budding at the posterior end, and is frequently combined with chain formation. The main characteristic is the entire absence of mouth and oesophagus, the animals being parasitic in the digestive tract of various annelids. Parasites, salt-water forms.
Anoplophrya branchiarum. Stein '52. Fig. 44.
_A. circulans_ Balbiani.
The body is cylindrical to pyriform, in the latter case broadened anteriorly. Cuticle distinctly marked by longitudinal striations which take the form of depressions and give to the body a characteristic melon shape. The endoplasm contains a number of large refringent granules--probably body products. The nucleus is elongate, somewhat curved, and coarsely granular. A micronucleus lies in the concavity. The cilia are long, inserted rather widely apart along the longitudinal markings. The contractile vacuole is single and is located at the pointed end, which is directed backwards during locomotion. One specimen found free swimming among some algæ.
Length 104µ; greatest diameter 36µ.
I was much surprised to find this form swimming about freely in the water; its mouthless condition showed it to belong to the family of parasites, the _Opalinidæ_. As the name indicates, however, this species is an ectoparasite upon the gills, and Stein gave the name _branchiarum_ to a fresh-water form parasitic upon _Gammarus pulex_. The Woods Hole form is so strikingly similar to the figure of _G. branchiarum_ that, although the name was given to a fresh-water form, it obviously applies to this marine variety. One important difference is the presence of only one contractile vacuole in the marine form.
KEY TO FAMILIES OF HETEROTRICHIDA.
Cilia cover the body 1
Cilia reduced to certain 2 localized areas
1. _Polytrichina_.
a. The mouth terminates a long Family _Plagiotomidæ_
peristomial furrow having an
adoral zone along the entire
left edge
b. Peristomial area a broad Family _Bursaridæ_
triangular area ending in mouth
c. Peristomial depression short; Family _Stentoridæ_
limited to the anterior end; its
plane at right angles to the
long axis of body; surface of
peristome striated and ciliated;
no undulating membranes
2. _Oligotrichina_.
a. Peristome without cilia; cilia Family _Halteriidæ_
limited to one or more girdles
about body
One marine genus *_Strombidium_
b. Thecate forms; the body is Family _Tintinnidæ_
attached by a stalk to the cup;
within the adoral zone is a ring
of cilia.
c. The peristomial depression is Family _Ophryoscolecidæ_
deep and funnel-like; cuticle
thick, with posterior spine-like
processes.
* Presence at Woods Hole indicated by asterisk.
KEY TO THE MARINE GENERA OF PLAGIOTOMIDÆ.
Diagnostic characters: The peristome is a narrow furrow which begins, as a rule, close to the anterior end and runs backward along the ventral side, to the mouth, which is usually placed between the middle of the body and the posterior end. A well-developed adoral zone stretches along the left side of the peristome, and is usually straight.
1. Body cylindrical; size medium; Genus _Metopus_
peristome long and turns sharply
to the left at the extremity
2. No torsion in the peristome; Genus _Blepharisma_
undulating membrane is confined
to the posterior part of peristome
3. No peristomial torsion; Genus _Spirostomum_
body highly contractile;
no undulating membrane
KEY TO THE MARINE GENERA OF BURSARIDÆ.
Diagnostic characters: The body is usually short and pocket-like, but may be elongate. The chief characteristic is the peristome, which is not a furrow, but a broad triangular area deeply insunk and ending in a point at the mouth. The adoral zone is usually confined to the left peristome edge, or it may cross over to the right anterior edge.
1. The anterior half of the body Genus _Balantidium_
tapers to nearly a point in front;
the peristome is narrowest at the
apex; the mouth is the entire
peristome base.
2. The anterior end does not taper; Genus *_Condylostoma_
the peristome is widest at the end
of the body; the mouth is clearly
defined.
* Presence at Woods Hole indicated by asterisk.
Genus CONDYLOSTOMA (KONDYLOSTOMA Bory de St. Vincent 1824) Dujardin '41
(Dujardin '41; Claparède & Lachmann '58; Stein '59, '67; Cohn '66; Quennerstedt '67; Wrzesniowski '70; Bütschli '76, '88; Kent '81; Maupas '83; Shevyakov '96.)
Colorless and more or less flexible animals of medium size. The general form is elongate and cylindrical or somewhat smaller anteriorly. The posterior end is broadly rounded, the anterior end somewhat truncate and oblique. The peristome is broad and triangular, the base of the triangle being the entire anterior end of the body. The entire length of the peristome is one-fourth or less of the body length. The mouth is large and placed at the apex of the peristomial triangle and opens into a comparatively small oesophagus. The right edge of the peristome is lamellate and bears a clearly defined undulating membrane. The adoral zone is well developed upon the left edge of the peristome, from which it passes around anteriorly to the right edge. The surface of the peristome is free from cilia, but the rest of the body is uniformly coated with small active cilia. Contractile vacuoles are not safely determined. Bütschli thinks there is probably one terminal vacuole, but some observers deny this (_e.g._ Maupas). Others describe them on the dorsal side of the posterior end (Quennerstedt). The macronucleus is long and beaded and placed upon the right side. Micronuclei are numerous and scattered along the macronucleus. The anus is terminal and dorsal. Food consists of large and small particles. Movement rapid, free swimming, alternating with resting periods; in some cases an undulating or wriggling movement is seen, showing clearly the flexibility of the body. Fresh and salt water.
Condylostoma patens Müller. Fig. 45.
The body is elongate, somewhat sac-like, five or six times as long as broad, plastic, and frequently contains brightly colored food granules. The triangular peristome takes up the greater part of the anterior end, and the mouth is situated at the sharper angle of the triangle, about one-fourth of the total length from the anterior end. The cuticle is longitudinally striated, the lines having a slightly spiral course. They are not closely set, and fine cilia are thickly inserted along their edges. The endoplasm is granular and viscous. The motile organs consist of an adoral zone of membranelles, which stretch along the left edge of the peristome and the front edge of the body. The right edge of the peristome supports an undulating membrane. The nucleus is moniliform and extends the full length of the left side; a number of micronuclei are distributed along its course (Maupas).
Length 400µ; diameter at widest part 105µ. Maupas gives the length from 305µ to 495µ; and Stein 376µ to 564µ. Very common.
For a more extended account of the structures, see the excellent description by Maupas '83.
KEY TO THE MARINE GENERA OF STENTORIDÆ.
Diagnostic characters: The peristome is relatively short and limited to the front end of the animal, so that its plane is nearly at right angles to that of the longitudinal axis of the body. The adoral zone of cilia either passes entirely around the peristome edge or ends at the right-hand edge. The surface of the peristome is spirally striated and provided with cilia. Undulating membranes are absent.
1. Peristome circular in outline; Genus _Stentor_
limited to the anterior end
2. The peristome is drawn out into two Genus _Folliculina_
wing-like processes; tube-dwelling
Genus STROMBIDIUM Cl. & Lach. '58.
(Stein '67; Bütschli '73; Fromentel '74; Kent '81; Gruber '84; Entz '84; Maupas '83. Bütschli '88.)
Small, colorless (except for ingested food) animals with characteristic springing movements. The form is usually constant, but in some cases may be plastic like _Astasia_; it is usually globular or conical, the posterior end being more or less pointed, the anterior end broadest. The latter is surrounded by a complete circle of the adoral zone, the oral end of which passes into a peristomial depression which extends deep into the middle of the body. The mouth, with a very small oesophagus, lies at the bottom of the inturned peristome. The region surrounded by the adoral zone is frequently drawn out into an anterior process, occasionally bearing a pigment mass. The ventral surface in some cases bears cilia, which may be distributed or restricted to a row of large cilia. Trichocysts are usually present and may be widely spread, limited to the posterior region, or arranged in a girdle about the middle. The contractile vacuole is simple, and posterior in position. The macronucleus is spherical and usually central in position. Movement is rapid swimming, combined with resting and floating periods, the latter usually terminated by a sudden leap.
Fresh and salt water; more common in the latter.
Strombidium caudatum Fromentel '74. Fig. 46, a, b, c.
Fromentel described a fresh-water form of this genus with a caudal appendage. The body is pyriform, broadly truncate on the anterior end, in the middle of which rises a papilliform process (Schnabel). On this process is a heap of pigment granules, which, however, are not constant. A ring of long cirri surround the anterior end and pass into the peristome, and from the left edge of this line of cirri a large adoral zone continues down to the mouth. The peristome is elongate and sac-form, and the mouth lies at the posterior extremity. With the exception of a caudal filament there are no other motile organs; this is about half as long as the body, structureless, hyaline, and sharply pointed. It splits up into a bundle of fine fibers upon treatment with caustic potash (c). The cirri emerge from minute hollows in the edge of the anterior border. The cortical plasm contains peculiar rod-like bodies, which look more like lines or markings than like rods or trichocysts. The nucleus is large, spherical, and placed in the center of the body. The contractile vacuole is posterior.
Length without appendage is about 35µ; greatest diameter 15 to 18µ. In decaying vegetable matter. Common.
Although Fromentel's species is incompletely described, it is very evident that the organism corresponds fairly well with the Woods Hole variety. His was a fresh-water type; this is marine, but the caudal filament and the contractile vacuole are similar. Certainly in this case the organism can not be regarded as a Vorticella broken off its stalk, as Kent '81 suspected. The anterior process with its pigment spot; the cirri, the spherical nucleus, the position of the vacuole, etc., are all opposed to such an interpretation which Kent applied to the original species. Neither can it be a Tintinnoid. I place it provisionally as _S. caudatum_.
KEY TO THE MARINE GENERA OF TINTINNIDÆ.
Diagnostic characters: Body attached by a stalk to a cup. Inside the zone of membranelles is a ring of cilia (par-oral).
1. The test is gelatinous and more or Genus _Tintinnidium_
less covered by foreign particles
2. The test is chitinous and clear. Genus _Tintinnus_
No foreign particles.
3. The test is chitinous; covered by Genus *_Tintinnopsis_
foreign particles, growth rings
frequent
4. The test is chitinous, often Genus _Codonella_
covered by foreign particles.
The test is marked by discoid,
circular, or hexagonal spots.
5. The test is perforated by pores Genus _Dictyocysta_
of circular or hexagonal form.
* Presence at Woods Hole indicated by asterisk.
Genus TINTINNOPSIS Stein '67.
(Stein '67; Kent '81; Daday '87; Bütschli '88.)
Medium-sized ciliates, inclosed in a chitinous lorica with embedded sand crystals. The form of the house, or lorica, varies greatly. In some cases the mouth opening is wide, giving the lorica a bell form; it may be long and tubular, short and spherical, or variously indented. The animal is attached, as in the closely allied genus _Tintinnus_, by a peduncle to the bottom of the lorica. The anterior end of the animal is inclosed by two complete circles of cilia; one, the outer, forming the adoral zone, is composed of thick tentacle-like membranelles, the other consists of shorter cilia within the adoral zone. The mouth leads into a curved oesophagus containing rows of downward-directed cilia (Daday). The entire body is covered with cilia, but as the lorica is always opaque these can be made out only when the animal is induced to leave the house. The only difference between this genus and _Tintinnus_ is the covering of foreign bodies--usually sand crystals. Movement is rapid and restless, and peculiarly vibratory, owing to the apparent awkwardness in moving the house. Salt water.
Tintinnopsis beroidea Stein, var. plagiostoma Daday. Fig. 47.
Synonym: _Codonella beroidea_ Entz '84.
The shell is colorless, thimble-shaped, with a broadly rounded posterior end. The body is cylindrical. The internal organs were not observed. Membranelles 24 in number. Length 50µ; greatest diameter 40µ.
Var. compressa Daday '87.
The posterior end of the shell is pointed, the lower third of the shell is swollen, the upper third is uniform in diameter and without oral inflation or depression. Nucleus not seen.
Length 70µ; greatest diameter 48µ.
Tintinnopsis davidoffi Daday. Fig. 48.
The shell is large, elongated, and provided with a considerable spine. The chitin of the shell is covered with silicious particles of diverse size. The internal structures were not observed.
Length of shell and spine 230µ; diameter of the oral aperture 54µ.
The variations of these species are considerable, and as the internal structures, such as the nucleus, are essential in fixing their systematic position, I place them as above, provisionally, and until further observations can be made.
KEY TO FAMILIES OF HYPOTRICHIDA.
a. Peristome indistinct; cilia on Family _Peritromidæ_
ventral surface uniform and not One genus, *_Peritromus_
differentiated into cirri
b. Peristome more or less indistinct; Family _Oxytrichidæ_
cilia reduced to a few rows on the
ventral surface; anal and frontal
cirri present
c. Cilia entirely reduced; frontal Family _Euplotidæ_
and anal cirri present or reduced;
macronucleus band-formed or spherical
d. Peristome reduced to left edge and Family _Aspidiscidæ_
does not reach over the anterior One genus, *_Aspidisca_
margin
* Presence at Woods Hole indicated by asterisk.
Genus PERITROMUS Stein '62.
(Stein '62, '67; Maupas '83.)
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Marine Protozoa from Woods HoleChapter II: Part 2
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