Chapter 1

E-text prepared by Ronald Calvin Huber, while serving as Penobscot Bay Watch, Rockland, Maine, and Joseph E. Loewenstein, M.D.

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Contributions from the Biological Laboratory of the U. S. Fish Commission, Woods Hole, Massachusetts.

MARINE PROTOZOA FROM WOODS HOLE.

by

GARY N. GALKINS, Department of Zoology, Columbia University.

_Bulletin of the United States Fish Commission_ 21:415-468, 1901

Comparatively little has been done in this country upon marine Protozoa. A few observations have been made by Kellicott, Stokes, and Peck, but these have not been at all complete. With the exception of Miss Stevens's excellent description of species of _Lichnophora_ I am aware of no single papers on individual forms. Peck ('93 and '95) clearly stated the economic position of marine Protozoa as sources of food, and I need not add to his arguments. It is of interest to know the actual species of various groups in any locality and to compare them with European forms. The present contribution is only the beginning of a series upon the marine Protozoa at Woods Hole, and the species here enumerated are those which were found with the algæ along the edge of the floating wharf in front of the Fish Commission building and within a space of about 20 feet. Many of them were observed in the water and algæ taken fresh from the sea; others were found only after the water had been allowed to stand for a few days in the laboratory. The tow-net was not used, the free surface Protozoa were not studied, nor was the dredge called into play. Both of these means of collecting promise excellent results, and at some future time I hope to take advantage of them.

My observations cover a period of two months, from the 1st of July to the 1st of September. During that time I was able to study and describe 72 species representing 55 genera, all from the limited space mentioned above. In addition to these there are a few genera and species upon which I have insufficient notes, and these I shall reserve until opportunity comes to study them further.

I take this opportunity to express my thanks to Dr. Hugh M. Smith for many favors shown me while at Woods Hole.

In dealing with these marine forms from the systematic standpoint, two courses are open to the investigator. He may make numerous new species based upon minor differences in structure, or he may extend previous descriptions until they are elastic enough to cover the variations. The great majority of marine protozoa have been described from European waters, and the descriptions are usually not elastic enough to embrace the forms found at Woods Hole. I have chosen, however, to hold to the conservative plan of systematic work, and to make as few new species as possible, extending the older descriptions to include the new forms.

The different classes of Protozoa, and orders within the classes, are distributed more or less in zones. Thus the Infusoria, including the Ciliata and the Suctoria, are usually littoral in their habitat, living upon the shore-dwelling, or attached, water plants and upon the animals frequenting them. It is to be expected, therefore, that in forms here considered there should be a preponderance of Infusoria. Flagellated forms are also found in similar localities, but on the Surface of the sea as well; hence the number described in these pages is probably only a small proportion of the total number of Mastigophora in this region. The Sarcodina, including the Foraminifera and the Radiolaria, are typically deep-sea forms and would not be represented by many types in the restricted locality examined at Woods Hole. Two species, _Gromia lagenoides_ and _Truncatulina lobatula_, alone represent the great order of Foraminifera, while the still larger group of Radiolaria is not represented at all.

The Protozoa described are distributed among the different orders as follows:*

Class SARCODINA. Subclass RHIZOPODA. Order AMOEBIDA. 1. _Amoeba guttula_ Duj 2. _Amoeba_ sp. 3. _Trichosphærium sieboldi_ Schn. Order RETICULARIIDA. Suborder IMPERFORINA. 4. _Gromia lagenoides_ Gruber. Suborder PERFORINA. 5. _Truncatulina lobatula_ Walker & Jacob. Subclass HELIOZOA. Order APHROTHORACIDA. 6. _Actinophrys sol_ Ehr. Order CHLAMYDOPHORIDA. 7. _Heterophrys myriapoda_ Archer. Class MASTIGOPHORA. Subclass FLAGELLIDIA. Order MONADIDA. 8. _Mastigamoeba simplex_, n. sp. 9. _Codonoeca gracilis_, n. sp. 10. _Monas_ sp. Order CHOANOFLAGELLIDA. 11. _Monosiga ovata_ S. Kent. 12. _Monosiga fusiformis_ S. Kent. 13. _Codonosiga botrytis_ (Ehr.) J. Cl. Order HETEROMASTIGIDA. 14. _Bodo globosus_ Stein. 15. _Bodo caudatus_ (Duj.) Stein. 16. _Oxyrrhis marina_ Duj. Order EUGLENIDA. 17. _Astasia contorta_ Duj. 18. _Anisonema vitrea_ Duj. Order SILICOFLAGELLIDA. 19. _Distephanus speculum_ Stöhr. Subclass DINOFLAGELLIDIA. Order ADINIDA. 20. _Exuviælla lima_ Clenk. 21. _Exuviælla marina_ Clenk. Order DINIFERIDA. 22. _Gymnodinium gracile_ Bergh. 23. _Glenodinium cinctum_ Ehr. 24. _Glenodinium compressa_, n. sp. 25. _Peridinium digitale_ Pouchet. 26. _Peridinium divergens_ Ehr. 27. _Ceratium tripos_ Nitsch. 28. _Ceratium fusus_ Ehr. 29. _Amphidinium operculatum_ Clap. & Lach. Class INFUSORIA. Subclass CILIATA. Order HOLOTRICHIDA. Family ENCHELINIDÆ. 30. _Lacrymaria lagenula_ Cl. & Lach. 31. _Lacrymaria coronata_ Cl. & Lach. 32. _Trachelocerca phoenicopterus_ Cohn. 33. _Tiarina fusus_ Cl. & Lach. 34. _Mesodinium cinctum_, n. sp. Family TRACHYLINIDÆ. 35. _Lionotus fasciola_ Ehr. 36. _Loxophyllum setigerum_ Quenn. Family CHLAMYDODONTIDÆ. 37. _Nassula microstoma_ Cohn. 38. _Chilodon cucullulus_ Müll. 39. _Dysteria lanceolata_ Cl. & Lach. Family CHILIFERIDÆ. 40. _Frontonia leucas_ Ehr. 41. _Colpidium colpoda_ Ehr. 42. _Uronema marina_ Duj. Family PLEURONEMIDÆ. 43. _Pleuronema chrysalis_ Ehr. 44. _Pleuronema setigera_, n. sp. 45. _Lembus infusionum_, n. sp. 46. _Lembus pusillus_ Quenn. Family OPALINIDÆ. 47. _Anoplophrya branchiarum_ Stein. Order HETEROTRICHIDA. Family BURSARIDÆ. 48. _Condylostoma patens_ Müll. Family HALTERIDÆ. 49. _Strombidium caudatum_ From. Family TINTINNIDÆ. 50. _Tintinnopsis beroidea_ Stein. 51. _Tintinnopsis davidoffi_ Daday. Order HYPOTRICHIDA. Family PERITROMIDÆ. 52. _Peritromus emmæ_ Stein. Family OXYTRICHIDÆ. 53. _Epiclintes radiosa_ Quenn. 54. _Amphisia kessleri_ Wrzes. Family EUPLOTIDÆ. 55. _Euplotes charon_ Ehr. 56. _Euplotes harpa_ Stein. 57. _Diophrys appendiculatus_ Stein. 58. _Uronychia setigera_, n. sp. 59. _Aspidisca hexeris_ Quenn. 60. _Aspidisca polystyla_ Stein. Order PERITRICHIDA. Family LICHNOPHORIDÆ. 61. _Lichnophora macfarlandi_ Stevens. Family VORTICELLIDÆ. 62. _Vorticella marina_ Greeff. 63. _Vorticella patellina_ Müller. 64. _Zoothamnium elegans_ D'Udek. 65. _Cothurnia crystallina_ Ehr. 66. _Cothurnia nodosa_ Cl. & Lach. 67. _Cothurnia imberbis_ Ehr. Subclass SUCTORIA. Family PODOPHRYIDÆ. 68. _Podophrya gracilis_, n. sp. 69. _Ephelota coronata_ Wright. Family ACINETIDÆ. 70. _Acineta divisa_ Fraip. 71. _Acineta tuberosa_ Ehr. Family DENDROSOMIDÆ. 72. _Trichophrya salparum_ Entz.

* This classification includes only the orders and families represented at Woods Hole

Genus AMOEBA Auct.

The pseudopodia are lobose, sometimes absent, the body then progressing by a flowing movement; the body consists of ectoplasm and endoplasm, the latter being granular and internal, the former hyaline and external. There is always one nucleus and one vacuole, but both may be more numerous. Reproduction takes place by division or by spore-formation. Fresh-water and marine.

Amoeba guttala Duj. Fig. 1.

A minute form without pseudopodial processes, extremely hyaline in appearance, and characterized by rapid flowing in one direction. The body is club-shape and moves with the swollen end in advance. A comparatively small number of large granules are found in the swollen portion, while the smaller posterior end is quite hyaline. Contractile vacuole absent, and a nucleus was not seen. Frequent in decomposing vegetable matter. Length 37µ. Traverses a distance of 160µ in one minute.

The fresh-water form of _A. guttula_ has a vacuole, otherwise Dujardin's description agrees perfectly with the Woods Hole forms.

Amoeba? Fig. 2.

A more sluggish form than the preceding, distinguished by its larger size, its dense granulation, and by short, rounded pseudopodia, which, as in _Amoeba proteus_, may come from any part of the body. A delicate layer of ectoplasm surrounds the granular endoplasm, and pseudopodia formation is eruptive, beginning with the accumulation of ectoplasm. Movement rapid, usually in one direction, but may be backwards or sideways, etc. Contractile vacuole absent; the nucleus is spherical and contains many large chromatin granules. Length 80µ; diameter 56µ.

Genus TRICHOSPHÆRIUM Schneider '78

Synonym: _Pachymyxa hystrix_ Gruber.

Marine rhizopods, globular or irregular in form, and slow to change shape. Dimorphic. Both forms multinucleate during vegetative life. Pseudopodia are long, thin, and thread-form, with rounded ends. Their function is neither food-getting nor locomotion, but probably tasting. The plasm of both forms is inclosed in a soft gelatinous membrane. In one form the jelly is impregnated with needles of magnesium carbonate (Schaudinn), but these are absent in the other form. The membrane is perforated by clearly defined and permanent holes for the exit of the pseudopodia. Reproduction occurs by division, by budding or by fragmentation, but the parts are invariably multinucleate. At the end of vegetative life the needle-bearing form fragments into numerous mononucleate parts; these develop into adults similar to the parent, but without the spines. At the end of its vegetative life this new individual fragments into biflagellated swarm-spores which may conjugate, reproducing the form with needles. Size up to 2 mm.

Trichosphærium sieboldi Schneider. Fig. 3.

With the characters of the genus. A form which I have taken to be a young stage of this interesting rhizopod is described as follows:

A minute, almost quiescent, form which changes its contour very slowly. The membrane is cap-like and extends over the dome-shaped body, fitting the latter closely. The endoplasm is granular and contains foreign food-bodies. Nucleus single, spherical, and centrally located. Pseudopodia short and finger-form, emerging from the edge of the mantle-opening and swaying slowly from side to side or quiescent. The most characteristic feature is the presence of a broad, creeping sole, membranous in nature and hyaline in appearance. This membrane is the only evidence of ectoplasm, and it frequently shows folds and wrinkles, while its contour slowly changes with movements of body. The pseudopodia emerge from the body between this membrane and the shell margin. Contractile vacuole absent. Length 42µ, width 35µ. In decomposing seaweeds, etc.

Only one specimen of this interesting form was seen, and I hesitate somewhat in placing it on such a meager basis. It is so peculiar, however, that attention should be called to it in the hope of getting further light upon its structure and mode of life. Its membranous disk recalls the genus _Plakopus_; its mononucleate condition, its membranous disk, and the short, sometimes branched, pseudopodia make it difficult to identify with any phase in the life-history of _Trichosphærium_. I shall leave it here provisionally, with the hope that it may be found more abundantly another time.

Genus GROMIA Dujardin '35.

(Dujardin 1835; M. Schultze '62; F. E. Schultze '74; Leidy '77; Bütschli '83; Gruber '84.)

The form is ovoid or globular, and the body is covered by a tightly fitting, plastic, chitin shell, which, in turn, is covered by a fine layer of protoplasm. The flexibility of the shell makes the form variable as in the amoeboid types. The thickness of the shell is quite variable. The pseudopodial opening is single and terminal. The pseudopodia are very fine, reticulate, granular, and sharply pointed, and form a loose network outside of the shell opening. Nucleus single or multiple. Contractile vacuole is usually absent. Fresh and salt water.

Gromia lagenoides Gruber '84. Fig. 4.

This species is not uncommon about Woods Hole, where it is found upon the branches of various types of algæ. The body is pyriform, with the shell opening at the larger end. The chitinous shell is hyaline and plastic to a slight extent, so that the body is capable of some change in shape. The shell is thin and turned inwards at the mouth-opening, forming a tube (seen in optical section in fig. 4) through which the protoplasm passes to the outside. The walls of this tube are thicker than the rest of the shell, and in optical section the effect is that of two hyaline bars extending into the body protoplasm. A thin layer of protoplasm surrounds the shell and fine, branching, pseudopodia are given off in every direction. The protoplasm becomes massed outside of the mouth-opening and from here a dense network of pseudopodia forms a trap for diatoms and smaller Protozoa. The nucleus is spherical and contains one or two large karyosomes. The protoplasm is densely and evenly granular, without regional differentiation. I have never observed an external layer of foreign particles, such as Gruber described in the original species.

Length of shell 245µ; largest diameter 125µ.

Genus TRUNCATULINA D'Orbigny.

A group of extremely variable foraminifera in which the shell is rotaline; i. e., involute on the lower side and revolute on the upper (Brady). The shell is calcareous and coarsely porous in older forms. The characters are very inconstant, and Brady gives up the attempt to distinguish the group by precise and constant characters.

Truncatulina lobatula Walker & Jacob.

Synonyms: See Brady '84 for a long list.

"It is impossible to define by any precise characters the morphological range of the present species. Its variations are infinite." (Brady, p. 660.)

This very common form, which occurs in all latitudes, was found frequently among the algæ at Woods Hole. Its characters are so difficult to define that for the present I shall limit my record to this brief notice. Size of shell 230µ by 270µ.

Genus ACTINOPHRYS Ehr.

The body is spherical and differentiated into granular endoplasm and vacuolated ectoplasm, but the zones are not definitely separated. There is one central nucleus and usually one contractile vacuole. The pseudopodia have axial filaments that can be traced to the periphery of the nucleus. Fresh and salt water.

Actinophrys sol Ehr., variety. Fig. 5.

Synonyms: See Schaudinn '95.

The diameter is about 50µ; the vacuolated ectoplasm passes gradually into the granular endoplasm. This is the characterization given _A. sol_ by Schaudinn, and it applies perfectly to the freshwater forms. If I am correct, however, in placing an _Actinophrys_-like form found at Woods Hole in this species, the description will have to be somewhat modified. In this form (fig. 5) there is no distinction between ectoplasm and endoplasm, and there is an entire absence of vacuoles. The nucleus is central, and axial filaments were not seen. The single specimen that I found looked much like a Suctorian of the genus _Sphærophrya_, but the absence of a firm cuticle and the presence of food-taking pseudopodia with granule-streaming makes it a very questionable Suctorian, and 1 place it here until further study throws more light upon it.

Diameter of body 40µ; length of pseudopodia 120 to 140µ.

Genus HETEROPHRYS Archer.

The body is globular with but slight differentiation into ectoplasm and endoplasm; one nucleus in the latter; contractile vacuoles one or many; pseudopodia on all sides, thin, and with peripheral granule-streaming; surrounded by a globular, rather thick coat of jelly, which is hyaline inside and granular on the periphery. Fresh and salt water.

Heterophrys myriapoda Archer. Fig. 6.

Synonym: _H. marina_ Hert. & Less. '74.

Diameter 25 to 80µ; pseudopodia twice as long as the body diameter; the plasm often contains chlorophyll bodies (Zoochlorella). The granular part of the gelatinous layer is thick (up to 10µ). The spine-like processes are very thin and short. (Schaudinn '95.) The marine form found at Woods Hole probably belongs to this species, as described by Schaudinn. The short pseudopodia which give to the periphery a fringed appearance are quite regularly placed in connection with the pseudopodia. The latter are not so long as twice the body diameter, the longest being not more than equal to the diameter of the sphere. The body inside of the gelatinous covering is thickly coated with bright yellow cells similar to those on Radiolaria. The animal moves slowly along with a rolling motion similar to that described by Pènard '90, in the case of _Acanthocystis_. Diameter of entire globe 35µ; of the body without the jelly 18µ. The extremely fine granular pseudopodia are 8 to 35µ long. Common among algæ.

This form was probably meant by Peck '95, when be figured "a heliozoön."

KEY TO ORDERS OF FLAGELLIDIA.

Small, body usually amoeboid; 1 or more Order MONADIDA. flagella; no mouth

Small; plasmic collar around the Order CHOANOFLAGELLIDA. flagellum

With 2 or more flagella; one trails Order HETEROMASTIGIDA. behind

With 3 or more flagella, none of which Order POLYMASTIGIDA. trails

Large; firm body wall; 1 or 2 flagella; Order EUGLENIDA. mouth or pharynx, or both

Medium size; with chlorophyll, Order PHYTOFLAGELLIDA. no mouth, usually colonial

Small; silicious skeleton; parasitic Order SILICOFLAGELLIDA. on Radiolaria or free (One genus, _Distephanus_ Stöhr)

KEY TO THE FAMILIES OF MONADIDA.

No mouth; 1 or 2 flagella: amoeboid Family _Rhizomastigidæ_ with lobose or ray-like pseudopodia

Mouth at base of single flagellum; Family _Cercomonadidæ_ plastic; no pseudopodia

One flagellum; inclosed in gelatinous Family _Codonoecidæ_ or membranous cups

One flagellum; tentacle like process Family _Bikoecidæ_ at base of flagellum; inclosed in cup

One main flagellum and 1 or 2 Family _Heteromonadidæ_ accessory flagella

KEY TO MARINE GENERA OF MONADIDA.*

Family _Rhizomastigidæ_:

1. Flagellum repeatedly thrown off Genus *_Mastigamoeba_ and reassumed in part

2. Flagellum never thrown off 3

3. a. Pseudopodia lobose Genus _Mastigamoeba_

b. Pseudopodia ray-like Genus _Mastigophrys_

Family _Codonoecidæ_:

1. Goblet-shaped cups adherent Genus *_Codonoeca_ by stalk

Family _Heteromonadidæ_:

1. The long flagellum vibratory Genus *_Monas_

2. The long flagellum rigid; Genus _Sterromonas_ shorter one vibrates

* Presence at Woods Hole indicated by asterisk.

KEY TO MARINE QENERA OF CHOANOFLAGELLIDA.

1. Without gelatinous 3 or membranous test

2. With gelatinous 4 or membranous test

3. a. Attached forms:

1. Without a stalk, or with Genus *_Monosiga_ a very short one

2. With a long, simple, stalk Genus *_Codonosiga_

3. With a long, branched, stalk Genus _Codonocladium_

b. Free-swimming Genus _Desmarella_

4. Colonial, and with a gelatinous Genus _Proterospongia_ covering

* Presence at Woods Hole indicated by asterisk.

KEY TO FAMILIES AND MARINE GENERA OF HETEROMASTIGIDA.

1. Two flagella nearly equal in size Family _Bodonidæ_

One main and 2 accessory flagella Family _Trimastigidæ_

Family _Bodonidæ_:

1. Body very plastic, Genus *_Bodo_ almost amoeboid

Body not plastic; with large Genus *_Oxyrrhis_ anterior cavity, holding flagella

Family _Trimastigidæ_:

1. With an undulatory membrane Genus _Trimastix_ between accessory flagella

Without such membrane; flagella Genus _Costia_ contained in a ventral groove while at rest

* Presence at Woods Hole indicated by asterisk.

KEY TO MARINE GENERA OF POLYMASTIGIDA.

1. Body flattened; ends rounded; Genus _Trepomonas_ sides hollowed; often with wing-like processes; cross section S-shaped

2. Body pyriform; one large Genus _Tetramitus_ asymmetrical groove; 4 flagella

3. Body spherical; many flagella Genus _Multicilia_ equally distributed

KEY TO FAMILIES AND MARINE GENERA OF EUGLENIDA.

1. With deeply-insunk pharynx; 2 no mouth

With pharynx and distinct mouth Family _Peranemidæ_

2. Body plastic; usually with Family _Euglenidæ_ chromatophores and eye-spot

Body plastic; no chromatophores; Family _Astastidæ_ no eye-spot

Family _Euglenidæ_:

Body _Euglena_-like, inclosed Genus _Trachelomonas_ in shell with round opening for exit of flagellum

Family _Astastidæ_:

Body with one flagellum Genus *_Astasia_

Family _Peranemidæ_:

1. Body striped; plastic; Genus _Heteronema_ two diverse flagella

2. Body striped; not plastic; Genus *_Anisonema_ posterior flagellum longer than the other

3. Body striped; not plastic; Genus _Entosiphon_ with rod-like organ in pharynx

* Presence at Woods Hole indicated by asterisk.

Genus MASTIGAMOEBA F. E. Schultze '75.

(Kent '81; Bütschli '86; Klebs '92; Senn 1900.)

In general the form is oval and either regular in outline or irregular through the presence of many pseudopodia. One flagellum usually quite large and distinct. Differentiation of ectoplasm and endoplasm distinct or wanting. One to several contractile vacuoles. The pseudopodia are occasionally withdrawn, and the flagellum is the sole means of locomotion. In some cases the flagellum turns into a pseudopodium, and, conversely, the pseudopodium at one end may become a flagellum (see below). In some rare cases the ectoplasm secretes a gelatinous mantle. Reproduction not observed.

Fresh and salt water.

Mastigamoeba simplex, n. sp. Fig. 7.