Part 1

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THE DIATOMACEÆ OF PHILADELPHIA AND VICINITY

BY

CHARLES S. BOYER, A.M., F.R.M.S.

_ILLUSTRATED WITH SEVEN HUNDRED DRAWINGS BY THE AUTHOR_

PRESS OF

J. B. LIPPINCOTT COMPANY

EAST WASHINGTON SQUARE PHILADELPHIA

1916

PREFACE

The present contribution to the local flora is intended as an introduction to more extended research.

The study is of advantage in relation to the life history of aquatic animals, the determination of ocean currents, as proved by polar discoveries, the investigation of geological strata where other fossil forms are absent, and the analysis of water supply; and, when we consider the universal distribution of diatomaceæ in the earth, the water and even in the air and the enormous deposits formed in past ages and still forming, we are able to realize the importance of a knowledge of these complicated forms and their function of purification.

The absence of descriptive works of reference in available form in this country, the polyglot confusion of authorities abroad and the amount of time, patience and skill required in obtaining, preparing and examining specimens, render the study one of difficulty.

The bibliography is omitted, as it is understood by those who possess the works of reference, and but few synonyms are given, having but little, except historical, value, especially when it is considered that modern investigators have no access to many of the earlier collections, when any of these exist.

So far as the marine forms are concerned, it is probable that nearly all occurring north of Florida are here included, and the fresh-water species described represent a large proportion of those found east of the Alleghanies. All of the figures are drawn to the same scale, a magnification of eight hundred diameters, from specimens in my possession, nearly all of which were found in or near Philadelphia.

If the work is of any value in inducing further investigation, I hope, in the words of Julien Deby, that "those who follow my advice will find in the study of these wonderful little organisms as much pleasure as I myself have found."

THE AUTHOR.

{5}INTRODUCTION

The Delaware River rises in the Western Catskill Mountains, flows southward for about three hundred and seventy-five miles, and expands into Delaware Bay about sixty miles from the sea. Its origin is among the Devonian and Carboniferous rocks, and in its course it passes through Silurian, Triassic and Cretaceous formations, finally reaching the Cambrian and Laurentian beds. It also drains regions of the glacial drift and beds which overlie overturned Miocene strata, and are sometimes mixed with them. From the mountains, nearly four thousand feet high, to the Bay, where the depth of water is not greater than seventy-five feet, the diatomaceous flora, from Alpine cascades to the salt marshes of New Jersey, contains a larger number of species than any other equal portion of the American coast.

The city of Philadelphia, about one hundred miles from the sea, lies at the junction of the Schuylkill with the Delaware, and much of the land near the rivers, especially southward, is flat and low, composed of recent alluvial deposits. In the central districts the ground is high, the deep sub-soil being mostly a dry gravel resting upon gneiss and schist, although it is in part composed of a bluish clay which was probably laid down in the bed of the ancient river before the last period of the glacial drift. The blue clay was not all deposited at the same time, as in the lower strata many marine forms are found which do not occur in the upper layers. This is notably the case in a deposit obtained at Spreckel's Sugar Refinery and also at the east end of Walnut Street Bridge, where a layer of blue clay occurs which is overlain by glacial drift. In other parts of the city mixtures of blue clay with more recent deposits are found, including fresh-water forms from numerous creeks and rivulets which traversed what is now the city proper, and especially from the vicinity of Fourth and Market Streets, where there existed as late as the year 1700 a large pond known as the "Duck Pond" which was subject to tidal overflow from its outlet, Dock Creek. The river water at Philadelphia is not noticeably brackish, although the tide extends thirty miles above the city and, before the building of Fairmount Dam, to the Falls of the Schuylkill. At certain times, when the river is low, the influx of tide water is sufficient to produce an abundance of brackish water diatoms at Greenwich Point. The entire absence, however, at present, of many of the marine forms obtained in dredgings in the Delaware opposite the city, as at Smith's Island, now removed, and in certain well borings at Pavonia, Pensauken, Gloucester and other places in New Jersey, where the depth reached the old blue clay, indicates conditions quite different from those now prevalent. In the Bay itself comparatively few living species are found, at least in any abundance.

In the study of local forms which follows, the district included may be considered as circumscribed by the circumference of a circle having a radius of one hundred miles from Philadelphia, containing the States of New Jersey and Delaware, the southeastern part of {6}Pennsylvania, a portion of Maryland on the south and extending eastward to New York Bay and Long Island Sound as far as New Rochelle.

The greater number of fresh-water species described have been obtained from near the city along the Darby, Crum, Ridley and Brandywine Creeks and from various places in New Jersey, including the Pine Barren region of the southern part of the State. Numerous collections have been made in the Schuylkill and the various reservoirs and along the Wissahickon, "where an Alpine gorge in miniature of singular loveliness is to be found within the limits of a city." The fossil deposits are from well borings near Camden, N. J., and from excavations in various parts of the city.

There appears to be no relation between the Miocene beds of the eastern coast and the deposits here described, all of which have been formed later than the glacial period or in an interval between two such periods. Apparently no diatoms grew during the glacial era, at least in sufficient abundance to leave any perceptible traces of their existence. An examination of glacial "flour" and clays from the Catskills shows an entire absence of these forms, and I have never found them in the milky flow from the glaciers of the Alps nor in the constantly muddy streams in certain of our Western States. The opacity of the water produces the same result as the absence of light in the deep lakes of New England, where diatoms are found only on the stalks or roots of water-plants near the shore, while in shallow ponds, such as the small lake near the summit of Mt. Lafayette, the growth is abundant. Certain species will grow wherever there are moisture, light and heat, but the greater number require the presence, in small amounts, of substances produced by the decay of animal and vegetable life. An abundance of diatoms in fresh water is usually an indication of its potability, while their entire absence in shallow water may be due to an excess of bacteria.

The specimens from which the drawings are made have been collected by the author for many years; in addition to possessing an almost complete library on the subject, he has had the advantage of examining material obtained by the late Mr. Lewis Woolman and numerous slides furnished by a number of friends, including Mr. John A. Shulze, Mr. Frank J. Keeley and Mr. T. Chalkley Palmer, to whom I here take pleasure in expressing my thanks.

The difficulties of the study are well stated by Agardh in the following extract from the preface to his Systema Algarum:

"Because, indeed, in this respect, no one will wonder whether in the distinction of species and reference to synonyms we have, perchance, committed many errors. They have occurred and are bound to occur, partly from the fact that one is not permitted to see the original specimens of all authors; partly, because sometimes even the original specimens of these plants are erroneous; partly, because the figures and descriptions of authors are often lacking and imperfect....

"There is added the difficulty of the study itself of these plants, their submerged habitat, the minuteness of their structure, the rarity of their fruit, the change in the dried {7}plant, the impossibility of culture, the fallacies of microscopical vision and the chaotic condition of Algology itself to-day."

The words of Agardh, written in 1824, are almost as true to-day. The lack of authentic specimens, which we hope will be remedied in time by the collections of the Smithsonian Institute, numerous incorrectly labelled slides in amateur collections, the imperfections of figures copied and recopied, without regard to relative size or correct references, and the confusion in the attempts to harmonize different descriptions, deter the student at the outset. The remaining difficulties mentioned by Agardh add, however, to the remarkable interest these forms have always had, since no increase in optical perfection of the microscope serves to lessen the mystery of their structure and mode of growth.

CLASSIFICATION

The few species of diatoms first discovered were included by Lyngbye, Dillwyn, and others in the genus _Conferva_. In 1824, the species, increased to forty-eight, were separated by Agardh into eight genera distinguished partly by their mode of growth. But little change was made until Heiberg, in 1863, advocated the division into symmetrical and asymmetrical forms. Without entering upon a general review of the later classifications, including Pfitzer's and Petit's divisions according to the number and location of the chromatophores, or the arrangement of Prof. H. L. Smith, because of the presence or absence of a raphe, or that of Mereschkowsky into motile and immotile forms, the modification of all of these methods by Schuett is here adopted, varied in accordance with certain monographs which appear to offer advantage.

It is customary, especially among writers who are familiar with other classes of plants, to decry any classification of diatoms according to the markings of their siliceous envelopes. As, however, one of the chief distinctions of the class is the possession of a more or less siliceous and indestructible frustule, and as the cell and its contents are never seen except within the valves, their variety forms the only available method of identification. The cell contents, owing to the difficulty of observing their living condition, their continued change, their lack of distinct variation and their entire absence in fossil forms, render their consideration as a complete method of classification an impossibility. If, however, the cell contents can be brought into relation with the markings of their siliceous envelope, it will be a consummation for which the future student of these complicated forms ought to be grateful. That this result is one to be expected may be inferred from the fact that the arrangement of protoplasmic masses in the interior of the cell is coincident in some cases with markings on the valve, and the character of the endochrome is assuming a certain value in accentuating the difference between such forms as _Pleurosigma_ and _Gyrosigma_, or in the resemblance between _Hantzschia_ and _Nitzschia_, or between _Surirella_ and _Campylodiscus_. Mereschkowsky, however, states that it is necessary to be careful in "establishing the relationship between diatoms based on the resemblance of their chromatophores," {8}and further observes that in _Hantzschia amphioxys_, _Scoliotropis latestriata_ and _Achnanthes brevipes_, three widely separated forms, the chromatophores are essentially the same.

In one of the earliest classifications of diatoms, the individual cell received less consideration than the nature of the filament or thallus in which many species occur in the first stages of their growth. Those, however, which exist in colonies at first are, sooner or later, broken up into separate frustules, either before or at the time of their maturity or previous to conjugation, while very many species are never seen except in a free state. The union of frustules, therefore, is of secondary importance and the group must be considered as filamentous or unicellular algæ. Their relation to other algæ is not well determined. Among the _Desmidiaceæ_, a family of the order _Conjugales_, of the class _Chlorophyceæ_, the cells are in many forms divided by a constriction into symmetrical halves. The Conjugales are starch forming, with walls of cellulose. In the Diatomaceæ the starch is replaced by oil globules, while the walls of cellulose are more or less filled with a deposit of silica. The Conjugales, however, reproduce by zygospores and usually contain pyrenoids, as may be seen in the parietal chromatophores of _Spirogyra_. In the class _Heterokontæ_ we have the reserve material in the form of oil, instead of starch, but there are no pyrenoids. To this class belongs the order _Confervaceæ_, in which the cells are unicellular or filamentous, and to which all of the Diatomaceæ were referred. While, therefore, Diatomaceæ have a close affinity to the Desmidiaceæ and to the Confervaceæ, the determination of their origin, one from another, or from a common ancestral type, appears to be a matter of conjecture.

{9}MORPHOLOGY AND DEVELOPMENT

THE CELL

The cell membrane is composed of two usually equal parts, each of which consists of a valve and a girdle or zone formed of cellulose modified by silica deposited in an insoluble state from a very dilute aqueous solution. The valves are more siliceous and robust than the girdle. Both are in most species easily separable, or at least the bands of the girdle which may be more or less closely fastened to the valves have a motion over each other permitting the cell to enlarge at pleasure. The longitudinal diameter of the cell, or the distance between the centres of the two valves, will vary according to the convexity of the valve and the age of the frustule which may be often determined by the width or number of the girdle bands. These, owing to their diversity of form and arrangement, will be further described under the generic diagnoses.

The siliceous cell-wall is covered on the outside by a layer of protoplasm called the coleoderm. This layer may be quite thin and evident only when treated with fuchsin or Bismarck brown, or it may be of considerable thickness. The cell contains the cytoplasma, protoplasm, cell-sap, endochrome, pyrenoids, oil globules and nucleus, together with certain other less understood bodies.

The Cytoplasma is a thin skin of colorless plasma covering the entire inner surface of the cell. It is invisible in the living cell but is evident in plasmolysis. In long forms it is thickened at the ends and is condensed at the plasma bridge which frequently connects the two valves and divides the cell into two parts, each containing more or less protoplasm surrounding the vacuole in which are found the cell-sap and certain granules. In some forms, as Meloseira, the cytoplasma includes the entire mass of protoplasm.

The Endochrome is seen in the form of one or more bands or plates, of a yellowish or brownish color, on the inner side of the valves or connective zone, or in granules or irregular masses, more or less numerous, on the inner walls, or sometimes grouped near the centre. It consists of a mixture of chlorophyll and diatomine which differ in their relative solubility in alcohol and in their spectroscopic analyses. The color varies from green to a chocolate brown in proportion to the amount of diatomine. So far as the function of the endochrome is concerned it does not appear to differ from that of ordinary chlorophyll, absorbing, under the influence of light, the carbon, and disengaging the oxygen of the carbonic anhydride in the water. Diatoms do not live in absolutely pure or non-aërated water. The individual plates or granules of the endochrome are called chromatophores. Their number and significance will be referred to in the description of genera.

THE PYRENOIDS.--In the chromatophores of many species are found colorless, homogeneous bodies, strongly refractive, of various shapes, usually lenticular or fusiform, which are known as Pyrenoids (Schmitz). They are scarcely evident in the living cell, but are distinguished by the action of hæmatoxylin and other reagents. Flat forms occur in Surirella and Pleurosigma, lens forms in Pinnularia, Stauroneis, Synedra, Fragilaria and Nitzschia, while a spherical form is found in Cymbella cuspidata. The pyrenoids are always imbedded in the chromatophore. Their growth is by division. Schmitz considers them a part of the living chromatophore, and their substance as working material which in excess has become resolved into the nature of a crystal which its form sometimes resembles. Comparisons are made between them and crystalloids found in certain monocotyledons. The pyrenoid is evidently concerned in the formation of the chromatophore, or in its division. Much of the conjecture, however, is due to the behavior of pyrenoids in other plants.

{10}OIL GLOBULES.--It has been established by Pfitzer that starch and sugar, as assimilation products, are replaced by oil in the cells of diatoms ("da bekannlich Staerke und Zucker bei den Bacillariaceen nicht nachzuweisen sind"). The oil drops are more or less numerous, of various sizes, and are found in the cytoplasma, the cell-sap, and sometimes the chromatophores. Mereschkowsky describes certain globules as elæoplasts, which he divides into four kinds according to their number and position. Whether all of these are oil globules is a question not yet determined.

Other bodies, known as "Buetschli granules," or volutin, and described as "little blisters filled with a tolerably robust refractive substance," are considered by Lauterborn to be a nitrogen reserve store. They are found in the cytoplasma, or in the cell-sap, and can be fixed in picric acid and stained in methylene blue.

NOTE.--For a discussion of the morphology of diatoms and a valuable résumé of the investigations of Buetschli, Karsten, Lauterborn, Mereschkowsky, Mueller, Pfitzer, Schuett, and others, the student is referred to "Der Bau der Diatomzelle," by Dr. Otto Heinzerling, in "Bibliotheca Botanica," 1908.

CELL DIVISION

The growth of diatoms follows the usual method of cell division as described by Sachs (Text Book of Botany, 2nd ed., p. 16): "The nucleus of a cell which is about to divide becomes broader, assuming the form of a biconcave lens, and its nucleolus breaks up into irregular granules which together with its other granular contents begin to form a nuclear disc in the equatorial plane. A delicate striation is now apparent in what is becoming the long axis of the nucleus, at right angles to the nuclear disc, and the characteristic nuclear spindle is gradually produced. The nuclear disc splits into two halves lying side by side, each of which travels to the corresponding pole of the nucleus; thus two nuclei are constituted which are connected by fibrillæ."

The cell-wall and the chromatophore bands divide, each nucleus passes to the centre, and two new cells are formed. In the meantime, to permit of this division, the two siliceous valves separate, the girdle bands slipping over each other, and opposite the larger or enclosing valve a new valve is formed, the girdle band of which is seen later within the girdle of the mother valve. Opposite the smaller valve of the original cell and adjoining the new valve, another valve is formed which also produces a girdle within the girdle of the smaller valve. As a result of division we have, therefore, the valves of the original, or mother cell, the two new valves and four girdle bands. (Pl. 40, Figs. 18 and 19.)

In the process of division, the continual formation of new valves, enclosed in the older girdle bands, will naturally cause a reduction in the size of the frustule. While this reduction, owing to the elasticity of the girdle, does not always occur, I believe, yet, in most cases, the diameter is so reduced that a rejuvenescence of growth is required. This is caused by the production of auxospores which may appear without conjugation. In this process, the beginning of which, in certain species, may be noticed by the increase in the size of the girdle as in reduplication, the two valves separate and within is formed a more or less spherical mass about twice the size of the original frustule and which forms on its circumference two large and often shapeless valves. These valves form others which assume the appearance of the original valves, but larger, and proceed to grow in the usual way. The reduction in size of the frustule seldom proceeds further than about half the size of the type form, so that, as a general rule, it may be stated that diatoms are not often smaller than half the larger size.

REPRODUCTION

The process of reproduction has been observed in many cases, but the conclusions reached are somewhat at variance with each other. The auxospore formation is simply a {11}method of rejuvenescence. When, however, the auxospores are thrown off from filamentous diatoms, it is probable that two may conjugate, their contents dividing each into two daughter cells which unite into two zygospores. The usual method is the union of two frustules, which, throwing off the old valves, coalesce into a single mass of protoplasm which produces an auxospore, sometimes called a sporangial frustule. It is stated that in some cases two frustules coalesce and produce two auxospores.

The existence of spores in diatoms is a much-disputed point. While they have never been seen, the inference that they exist is very great, as otherwise it becomes difficult to understand the sudden growth of species in localities and under conditions that seem to preclude the actual presence of the living frustule. It is a matter of common observation that, in examining collections of living forms, minute frustules or brownish globules appear to resemble larger diatoms. In gatherings of Gomphonema, when many specimens are sessile on the same object, numerous intermediate sizes, varying from minute globules to the type, are seen, yet not positively demonstrable as the same.

Conjugation, the formation of auxospores, and the actual process of cell division are seldom seen, as they occur during the night or at least in darkness. It is advisable in order to observe reduplication to obtain the material about midnight and place it in very dilute alcohol. In filamentous forms, however, the cell division is easily observed at any time in its various stages. By immersing in picric acid (saturated solution), transferring to very dilute alcohol which is gradually increased in strength, and then passing through oil of cloves and finally to the mounting medium, excellent preparations can be made. By staining with gold chloride alone the nucleus is made apparent without further treatment.

EVOLUTION OF FORMS