Diatomaceae of Philadelphia/Introduction
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 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 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," 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.