TbeVcqe'tation o{ ihe •1 ME •lilvlrls^vl^ ri)cr-r ^,^1 y^^^ \ I Rutgers University Libraries ^7 GIFT OF MARIE A. RICHARDS Digitized by the Internet Archive in 2010 with funding from Lyrasis IVIembers and Sloan Foundation http://www.archive.org/details/vegetationofnewjOOhars Figure i Cedar swamp at Shamong, June 27, 1910, with an even stand of Chamaecyparis thyoides and a sphagnum basis out of which grow Rhus vernix, Acer rubrum, Ciethra alnifolia, Myrica caroiinensis, Vaccinium corymbosum. Note the fronds of Osmunda cinnamomea and the low herb, Trientalis americana. (Photograph by Henry Troth and J. W. Harshberger.) THE VEGETATION OF THE NEW JERSEY PINE-BARRENS AN ECOLOGIC INVESTIGATION BY JOHN W. HARSHBERGER, Ph.D. PROFESSOR OF BOTANY, UNIVERSITY OF PENNSYLVANIA MEMBER OF THE AMERICAN PHILOSOPHICAL SOCIETY AND OF THE ECOLOGICAL SOCIETY OF AMERICA, FELLOW OF THE AMERICAN GEOGRAPHICAL SOCIETY OF NEW YORK, ETC. PHILADELPHIA CHRISTOPHER SOWER COMPANY 124 NORTH EIGHTEENTH STREET 1916 Copyright, 19 i6, by John W. Harshberger WM. F. FELL CO. PRINTERS PHILADELPHIA FOREWORD The following pages will be devoted to a description of the pine-barren vegetation of the coastal plain of New Jersey from the phytogeographic and the ecologic aspects. The bulk of the monograph will present the observations and the research work which have been in progress for a period of at least twenty-five years, although the greater part of the material has been collected during the last ten years. Besides the original material, which comprises the larger part of the monograph, emphasis will be laid upon what has been published on the subject as it elucidates, or illumines, the problems which have been kept in view during the prosecution of this botanic study. The thought is to collate all the facts which are not purely statistic and systematic and which bear in any way upon the pine-barren region of New Jersey, or upon its interesting and characteristic vegetation. Full details of a general historic, statistic, tloristic, and systematic character will be found in Witmer Stone's volume, issued in 1 91 1 by the New Jersey State Museum, upon "The Plants of Southern New Jersey with Especial Reference to the Flora of the Pine Barrens and the Geographic Distribution of the Species."* Many facts which the writer intended originally to incor- porate in this monograph have appeared in the work of Stone, so that attention will be given chiefly to matters which concern the plant geog- rapher, plant ecologist, and plant physiologist. As an almost complete bibliography of the flora is given by Stone, such a list will be omitted entirely, as will also facts which bear upon the distribution of plants in southern New Jersey as related to that of other parts of our country. Although the material for this work has been gathered contemporane- ously with that of Stone and our confreres in the Philadelphia Botanical Club, but entirely independent of them, it will be made to supplement that volume by including only that which has not been mentioned by Stone, or in a very casual and unemphatic way. The illustrations in the Annual Report of the New Jersey State Museum largely depict the species of plants which are characteristic of southern New Jersey and the pine-barrens, and a few that show actual conditions of vegetation. * Stone, Witmer: The Plants of Southern New Jersey, with Especial Reference to the Flora of the Pine-Barrens and the Geographic Distribution of the Species. Annual Re- port of the New Jersey State Museum, 1910, pp. 25-828, with 129 plates; Trenton, 191 1. VI VEGETATION OF THE NEW JERSEY PINE-BARRENS The illustrations of this monograph will show the natural vegetation of the region as it appeals to the plant ecologist, landscape artist, and nature lover. The maps and other figures, therefore, will be of an entirely different character from those given in the bulky volume of the New Jersey State Museum. The demand for such a work as this one following so closely upon that of the museum report has been emphasized by Harper* in his review of Stone's book, where he states that "with such a splendid floristic founda- tion to build on, the time is now ripe for some ecologically inclined botanist to make a detailed study of the vegetation of the same region, and thereby fill a long-felt want." Such a study is pressing, because the original conditions of the coun- try are fast changing. Forest fires have done great damage to the wild plants. The spread of areas of cultivation has destroyed many tracts of unusual botanic interest. The construction of dams, of railroad embankments and bridges across streams has wrought decided changes in the original pine-barren vegetation. The following account, there- fore, will be a printed record, embellished with photographs and other illustrations, of the pine-barren vegetation in its original, wild con- dition, as far as that could be determined at the present time. The author wishes to thank with deep appreciation the financial as- sistance given by the following persons, without whose aid the publica- tion of the book would have been impossible. American Philosophical Society. Henry D. Moore. Edward Bok. H. K. Mulford. Henry G. Bryant. Clement B. Newbold. Herman Burgin. Sarah Nicholson. Sarah C. DeHaven. Harold Peirce. Frank R. Ford. James L. Pennypacker. William H. Greene. John T. Pennypacker. Joseph R. Grundy. Richard A. F. Penrose. Charles C. Harrison. W. Hinckie Smith. Samuel Heilner. J. W. Sparks. A. J. Hemphill. Thomas W. Synnott. Alba B. Johnson. Joseph H. Taulane. Adolph W. Miller. J. B. Van Sciver. He desires to acknowledge the encouragement of Harold Peirce and James L. Pennypacker in the publication of the book, and Roland M. Harper and James L. Pennypacker, who gave the galley proofs a careful reading. Miss Elizabeth Caley, a graduate of the Pennsylvania School * Harper, Roland M.: Stone's Flora of Southern New Jersey. Torreya, 12: 216- 225, September, 1912. VEGETATION OF THE NEW JERSEY PINE-BARRENS vii of Industrial Art, assisted in getting many of the drawings into final shape from pencil sketches furnished her by the author. Mrs. S. B. Moulton made the sketch of Pyxidanthera barbulata for the front cover of the book. The shade of green used in the cloth cover corresponds with that of the evergreen foliage of the pitch-pines as illumined by the sun on a bright summer day, the effect being heightened by the roughened sur- face of the cloth. The back-bone design represents the grass-fern, Schizaea pusilla, and the front-cover design, a spray of the flowering- moss, Pyxidanthera barbulata, both characteristic plants of the pine- barrens of New Jersey. Through the interest of Alfred Gaskill, Chief of the Division of Fores- try and Parks, Department of Conservation and Development of New Jersey, permission was given by the New Jersey Geological Survey to use its map of 191 5 as a base map upon which the details of the pine- barren vegetation have been printed in color. This has been done as faithfully as the constant shifting of the relative positions of the wild and cultivated areas of the region has permitted, as also the changes produced by fires and the axes of lumbermen, especially in the removal of the valuable white cedar trees from the swamps of the southern part of the state. It is noteworthy that a white cedar swamp of a month ago may become a second growth deciduous swamp of the following year. Such changes are not always represented on a map based upon surveys made before such changes take place. Philadelphia, October 5, 1916. TABLE OF CONTENTS CHAPTER 1 PAGES Physiography of the Region 1-7 CHAPTER H Geography of the Pine-Barren Region 8-11 Origin of tiie Designation "Pine-Barren," 8; Geographic Place Names, 9 CHAPTER HI Influence of Geography and Vegetation on the Peoples and the Indus- tries OF the Region 12-30 General Considerations, 12; Floral Districts, 15; Local Plant Names, 16; Industries of the Pine-Barrens, 17; Lumbering, 17; Turpentine Industry, 2 i ; Cranberry Culture, 25; Sphagnum, 26; Peat, 26; Huckleberry Picking, 27; Drugs, 27; Greens and Flowers, 29 CHAPTER IV Research Investigation of Pine-Barren Soils 3 1-47 General, 31; Mechanic Analysis, 33; Mineralogic Analysis, 34; Chemic Analysis, 34; Experiments with Soils, 35; Capillary Rise of Water, 42; Re- tention of Water, 43; Wilting Coefficient, 43; Moisture Equivalent, 47 CHAPTER V Phytogeographic Formations 48-86 Classification, 48; Pine-Barren Formation, 49; Architectural Forms of Pine Trees, 53; Facies of Pine-Barren Formation, 56; Method of Investigation, 57; High Pine-Barrens, 60; Flat Pine-Barrens, 65; Low, or Wet Pine-Bar- rens, 75; Colors of Pine-Barren Vegetation (Winter Aspect, Spring Aspect, Summer Aspect, Autumn Aspect), 77; Coastal Pine-Barrens, 82; Contrast of German and New Jersey Formations, 85 CHAPTER VI Pine-Barrens Transitional to Sea-Dune Vegetation 87-104 Transition from Sea-Dunes to Pine Thicket at Sea Girt, 90; Bluff Forest along Great Egg Harbor Bay at Somers Point, 91 ; Tension Line Salt Marsh to Pine Forest at Somers Point, 94; Transition Salt Marsh to Pine Forest between Ocean Gate and Barnegat Pier, 95; Pine-Barrens at Northern Limit, 96; Transition Forest Western Pine-Barren Limit, 97; Notes on the Distribution of Transition Plants, 100; Successional Formations (Natural, Oak-Coppice Succession in Transition Region, Oak-Coppice Succession in Cape May Region — Mixed Pine-Oak Succession — Coastal Oak Forest Suc- cession— Oak-Bottoms), 100-104 CHAPTER Vll Pine-Barren Plants as Weeds 103-1 13 Introduced Weeds, 105; Classification of Alien Plants, 106; Pine-Barrens of Long Island, 107; Tension Line between Pine Forest and Cedar Swamp, 109 ix X VEGETATION OF THE NEW JERSEY PINE-BARRENS CHAPTER VIII PAGES Cedar Swamp Formation 1 14-13 1 General, 114; Synecology of Cedar Swamps, 120; Additional White-Cedar Swamp and Bog Plants, 126; Seasonal Aspects, 127; Fungous Diseases of the White Cedar, 129; Additional Fungi Found on White Cedar, 131 CHAPTER IX Transition Pine Forest to Deciduous Swamp 132-142 General, 132; Deciduous Swamp Formation, 133; Branch Swamps, 136; Deciduous Swamps of Transition Region, 136; Reed Marsh Formation, 138; Additional Swamp Plants, 138; Stream Bank Formation, 139 CHAPTER X Pond Plant Formation 143-1 50 General Consideration, 143; Additional Plants, 144; Savanna Formation, 145 CHAPTER XI Plain Formation (Coremal) 151-171 Geographic Location, 151; Synecology, 154; Measurement of Species, 159; Theories Concerning Coremal, 159; Experimental Treatment of Soils, 162; Contrast with Nantucket and European Conditions, 166; Hempstead Plains, Long Island, 170 CHAPTER XII Cultivated Plants of the Pine-Barren Region 172-174 Forest Trees, 172; Fruit Trees, 172; Bush Fruits and Vines, 173; Small Fruits, 173; Root Crops, 173; Leaf Crops, 173; Fodder and Cereal Crops, 173; Flower Crops, 173; Garden Crops, 174 CHAPTER XIII General Observations on Pine-Barren Vegetation 175-184 General, 173; Pine-Barren Vegetation by Quadrats, 176; Generic Coefficient, 181; Families of Pine-Barren Plants, 182 CHAPTER XIV Biologic Types of Pine-Barren Plants 185-192 Raunkiaer's Classification, 185; Pine-Barren Formation (Trees, Shrubs, Undershrubs, Annual Herbs, Colorless Parasitic Plants, Ferns, Perennial Herbs, Evergreen Species, Deciduous-leaved Species), 186; Colors of Pine- Barren Flowers, 188; (do.) White-Cedar Swamp Formation, 190; (do.) Marshes, 192 CHAPTER XV Phytophenology of Pine-Barren Vegetation 193-215 Tables of Flowering and Fruiting Periods, 196-208 CHAPTER XVI Vegetative Propagation and Structure of the Shoots 216-244 Hess Classification, 217; Detailed Root and Stem Studies, 219-244 CHAPTER XVII General Remarks on Root Distribution 245-251 VEGETATION OF THE NEW JERSEY PINE-BARRENS XI CHAPTER XVI 1 1 PAGES Leaf Forms of Pine-Barren Plants 252-258 Classification, 253; General Remarks, 258 CHAPTER XIX Microscopic Leaf Structure 259-285 General, 259; Classification, 260; Detailed Structure, 263; Synopsis, 283 CHAPTER XX Cone and Seed Production of the Pitch-Pine 286-294 General Investigation, 286; Law of Cone and Seed Production, 292; Vivi- pary in the Acorns of Quercus marylandica, 292 CHAPTER XXI Notes on a Few Insect Galls 295-297 CHAPTER XXII Pine-Barren Plants from an Evolutionary Viewpoint 298-3 16 General, 298; Elementary Species, 299; Variation, 300; Mutations, 303; Epharmony. 305; Plasticity of Species, 305; Response to Ecologic Factors (Soil, Light, Wind, Water), 305; Convergent Epharmony, 308; Persistent Juvenile Forms, 312; Hybridization, 313; Conclusion, 313 Index 3 17-329 THE PINE-BARREN VEGETATION OF NEW |ERSEY CHAPTER I PHYSIOGRAPHY OF THE REGION The physiographic history of the coastal plain of southern New Jersey, which includes that portion of the state south and southeast of the "fall line," begins with the depression of land beneath the surface of the sea during the Lafayette epoch of the Miocene (Neocene) known as the Lafayette depression, or Miocene sinking. The Lafayette deposits were later elevated along the Atlantic and the Gulf Coasts, constituting in New Jersey the Beacon Hill formation and the older portion of the coastal plain land surface east and south of the "fall line." The materials deposited and later elevated above the sea consisted of gravel, sand, silt, and clay related variously to one another. This Post-Miocene period of uplift was associated with active erosion. The level plain which was developed as a result of the erosion of the Lafayette formation (the Beacon Hill of New Jersey) has been called the Pre-Pensauken peneplain in New Jersey.* This marginal plain extended along the Atlantic coast and was tenanted by a flora of fairly uniform character from north to south, for it is a noteworthy fact that out of a total of 386 New Jersey pine-barren plants. 183, or 48 per cent., represent an element common to the New Jersey and the southern pine-barrens, while 153 species are wide-ranging. This ancient Miocene coastal flora consists essentially of American generic and specific types. With the beginning of the Pleistocene, or early Glacial Period, the coastal plains of New Jersey, Maryland, and other Atlantic states were depressed in part and differently beneath the ocean in the submergence generally known as the Columbia, or Pensauken by New Jersey geolo- gists. The Beacon Hill (Lafayette) formation, however, remained con- stantly above water during the Pensauken submergence, constituting an * In the correlation of these formations I have consulted Chamberlain, T. C, and Salisbury, R. D.: Geology, in: 450; McGee, W.J.: The Lafayette Formation. Twelfth Annual Report, U. S. Geological Survey, 353-52 1, 1892. I I 2 VEGETATION OF THE NEW JERSEY PINE-BARRENS island tenanted by the ancient vegetation of the Miocene coastal plain. The vegetation of this New Jersey Pensauken island had reached already a climax condition when it was isolated by the Pensauken submergence. Taylor* in a recent paper has elaborated on the theory which the writer first presented in detail on page 220 of his Phytogeographic Survey of North America, where it is stated that "during the Pensauken submer- gence of the New Jersey geologists New Jersey was depressed to such an extent as to drown the Delaware River at its lower end, allowing the sea to pass up its valley and over the peneplain which had been developed during the previous cycle of erosion, so that a broad sound was formed which connected Raritan and Delaware bays, forming an island covered perhaps with pine-barren vegetation. The mouth of the Delaware* River during the Post-Pensauken uplift was transferred to Delaware Bay, followed by a cycle of erosion. It was during the Post-Pensauken uplift that the flora of the lower Delaware Valley and the coastal strip was probably developed, so that the New Jersey pine-barrens became sur- rounded by a fringe of vegetation developed along similar lines in the coastal strip and along the east and west banks of the Delaware River." This isolated island of pine-barren plants was removed still further from contact with the southern pine-barrens by the unequal depression of the coastal plain, so that with the exception of the island vegetation, the typic coastal plain plants were exterminated in the depressed portion of the plain in Delaware and Maryland during the Pensauken (the Wico- mico of Maryland) submergence, for Shrevef has indicated the almost utter lack of pine-barren plants in Maryland. Thus, if we contrast the area occupied by the Beacon Hill formation and the area in central New jersey covered by the pine-barrens, we find an unusual coincidence (Map, Fig. 2). This coincidence of the two areas is made clear when we remember that the Beacon Hill formation was an island during the Pen- sauken submergence and that it was covered with vegetation of the pine- barren type. The location of the present pine-barren area is due to this fact, to the present soil conditions, and to the fact that when the land surface was again elevated after the Pensauken submergence the decid- uous types of vegetation invaded the recently elevated land in the coastal plains of Delaware, Maryland, and New Jersey, so that the pine-barren vegetation was surrounded on all sides by deciduous forests. The decid- uous forests, on the other hand, were unable to supplant the pine-barren * Taylor, Norman: The Origin and Present Distribution of the Pine-Barrens of New Jersey. Torreya, 12: 235, October, 1912; Flora of the Vicinity of New York. Memoirs of the New Yori< Botanical Garden, v: 8-13, with suggestive maps, 1915. t Shreve, Forrest: Maryland Weather Service, hi: 87(1910). VEGETATION OF THE NEW JERSEY PINE-BARRENS Figure 2 Map showing geologic history of pine-barren region + + + Outer limit of Pre-Pensauken Peneplain. — . — . Inner limit (First Mountain) of Pensauken submergence. Dotted area under water. Boundary of Beacon Hill area. Not under water since the Miocene. Present boundary of pine-barrens. Note the overlapping of the Beacon Hill area and the pine-barren region. .... Great terminal moraine. Southern limit of glacial ice. A A. Rocky Hill. H H. Atlantic Highlands. 4 VEGETATION OF THE NEW JERSEY PINE-BARRENS vegetation, because the latter had reached a climax condition. Both types of vegetation in the New Jersey coastal plain were mutually ex- clusive. While the above outline was prepared before the appearance of Taylor's paper, yet in the elaboration of the theory by Taylor many important points are emphasized. To quote: " For the phytogeographer the salient features of these changes are that the Beacon Hill has been uninterruptedly out of water since AJpper Miocene times, and that it has several times been partly and often entirely surrounded by water. These facts, together with the encroachment of the glacier, and its re- cession, with the probable deposition of a great deal of morainic material around Beacon Hill, make this formation the oldest in New Jersey, either on the coastal plain or in the glaciated regions northward, that could have been continuously covered with vegetation. This, it seems to me, is why the Beacon Hill formation is the controlling factor in the origin and present distribution of the pine-barrens. The area of the pine-barrens (see Fig. 2) is not exactly coextensive with Beacon Hill, but the differences are so slight that recent and local erosion of the for- mation would account for the failure of the two regions to superim- pose, as it were." " In other words, the New Jersey pine-barrens exist exclusively on this Beacon Hill formation, an area isolated by geological processes, and maintaining a relict or climax flora, the antiquity of which greatly ante- dates any of the rest of our vegetation hereabouts, so far as permanency of position and phytogeographical isolation are concerned. This un- doubtedly accounts for the composition of the flora, and it is interesting to note that zoologists have found this same apparent isolation, the same endemism noted above." All of the facts presented above seem in- contestably to point to a geologic explanation of the origin and present distribution of the pine-barren vegetation.* An interesting light is thrown upon the origin of the pine-barren vegetation on an island land mass by contrasting the vegetation of the Island of Nantucket, where a low heath vegetation, representing many of the species in the under- growth of the New Jersey pine-barrens, is being invaded by the pitch-pine, so that some areas on Nantucket have become like the New Jersey pine- barrens in physiognomy and in floral constitution, illustrating the conver- sion of a treeless, wind-swept heath into a pine-heath, or Kiefernheide.f * The writer thinks it unfortunate that Taylor should make only a partial quotation of the subject matter found on pages 218 and 219 of the Phytogeography of North America, ignoring the views presented at the top of page 220 of the same book. The whole context should have been considered. t Harshberger, J. W.: The Vegetation of Nantucket. Bulletin Geographical Society of Philadelphia, xii : 24-33, Apr., 1914. Also consult the section of this monograph on the vegetation of the New Jersey plains, where additional evidence is presented. VEGETATION OF THE NEW JERSEY PINE-BARRENS 5 The glaciation of the northern part of North America had very Httle influence on the pine-barren region, for we find no evidence of the deposi- tion of glacial material, or of glacial terraces, but the surrounding region was not so fortunate, for the Post-Pensauken uplift inaugurated another period of erosion which was followed by a depression which allowed the deposit of materials that constitute the Cape May (Talbot of Maryland) subdivision of the Columbia. This effected only a small area of the state in early post-Glacial times. If we consider that during the uplift in post-Glacial times the coastal plain extended northeastward, tying together Long Island, Block Island, Martha's Vineyard, Nantucket, Cape Cod, and other New England islands as far as Newfoundland, we have a ready explanation of the presence of many coastal plain plants in all of these islands, and in Nova Scotia and in Newfoundland. There are 1 16 species of the New Jersey coastal plain that range from Florida to Massachusetts.* Fernaldf has shown that there is a southern coastal element in the flora of Newfound- land, especially in the sandy areas, or in the bogs on the carboniferous sandstones. These plants are of Nova Scotia, Sable Island, southern New England, Cape Cod, Long Island, New Jersey, and even farther south, such as: Panicum implicatum, Scirpus subterminalis, Carex silicea, C. hormathodes, C. sterilis, C. trisperma var. Billingsii, Eriocau- lon septangulare, Juncus pelocarpus, Habenaria blephariglottis, Spergu- laria rubra, Arenaria peploides var. robusta, Pyrus arbutifolia var. atropurpurea, Elatine americana, Hudsonia ericoides, Myriophyllum tenellum, Vaccinium macrocarpon, Gaylussacia dumosa var. Bigelowi- ana, Utricularia clandestina, Solidago uniligulata. Aster radula, A. nemoralis, Potamogeton Oakesianus, Rhynchospora fusca, Schizaea pusilla, and Corema Conradii. This class of southwestern Carolinian types includes 60 species, or 7^ per cent, of the Newfoundland flora. The distribution of Corema Conradii and Schizaea pusilla has long puzzled botanic geographers, but if we consider that in post-Glacial times the coastal plain continued far to the northeastward, in all prob- ability these two plants, as well as others, had a continuous distribution from south to north. In general, the areas covered by the ice of the Glacial Period have risen since the ice melted, but the New England coast has been depressed, unequally separating the coastal plain into a number of islands. Now the distribution of Corema Conradii tallies with this formation of islands by depression of the coast, for it occurs on * Stone, Witmer: Loc. cit. t Fernald, M. L.: A Botanical Expedition to Newfoundland and Southern Labrador. Rhodora, 13: 135, July, 191 1. 6 VEGETATION OF THE NEW JERSEY PINE-BARRENS the plains of the pine-barren "island" of New Jersey, on Long Island (on authority of Dr. Torrey), on Nantucket (abundant in 191 5), at Ply- mouth, Mass. (Tuckermann), Truro, Cape Cod (Watson), Bath, Maine (Gambell), Isle au Haut, Maine (Young), Kimball's Island, Maine, Mt. Desert, Maine, in Nova Scotia and Newfoundland.* Presenting these facts in sequence we have: (i) The wide dispersal of coastal plain plants on the Lafayette deposits during the Post-Mio- cene; (2) the isolation of the pine-barren vegetation on Pensauken Island during the Pensauken submergence which destroyed the coastal plain vegetation of Delaware and the eastern shore of Maryland in late Glacial time; (3) the introduction of the deciduous type of vegetation into the middle district of New Jersey, into Delaware and the eastern shore of Maryland during the Pensauken uplift of Interglacial time; (4) the spread of coastal plain plants northeastward to Nova Scotia and Newfoundland during the post-Glacial uplift; (5) the isolation and sepa- ration of coastal plain species, such as Corema Conradii and Schizaea pusilla, by the unequal depression of the New England coast with the formation of islands upon which many of these species have been pre- served, while in the depressed areas they have suffered extinction. Summarized in the form of a table, we have: Post-Miocene Time Wide dispersal of coastal plain plants. r-.„,„ /-, , T f The isolation of coastal plain plants on Pen- Larly Glacial 1 ime 1 i 1 j f k I sauken Is and. Inter-Glacial Time Post-Glacial Time The introduction of deciduous vegetation into the coastal plain of New Jersey. A. The spread of coastal plain plants north- ward with the extension of the land areas. B. The isolation of many such plants on coastal islands with later submergence. The coastal plain of New Jersey, as it stands to-day, does not in its highest point reach an altitude of 121 meters (400 feet) and it descends to sea-level about the borders of the state. It is a relatively flat plain with slight relief, but of a rolling character, with hills here and there, and shallow valleys occupied by the master streams of the country. The ocean-directed streams are longer and more frequently branched than the shorter Delaware-directed streams. The pine-barren region is traversed almost entirely by the rivulets, creeks and rivers connected with the Atlantic Ocean drainage system. Such are the Manasquan, * Redfield, John H.: Corema Conradii and its Localities. Bulletin Torrey Botanical Club.xi: 97-100; see also Plate 90, Figs. 9 and 10, of Fernald's paper. VEGETATION OF THE NEW JERSEY PINE-BARRENS 7 Toms, Wading, MuUica and Great Egg Harbor rivers. (Colored map, Plate I.) The physiography of the country, as reflected in the areas of lowland and highland controlled by the streams, has important influence on the location of the important plant formations of the region. The salient features of this relationship will be emphasized in subsequent pages. CHAPTER II GEOGRAPHY OF THE PINE-BARREN REGION The northern edge of the pine-barren region of coastal New Jersey is in approximately 40° 15' N. latitude. The southern edge is found at 39° 10' N. latitude. The extreme western border is at Maurice River, at latitude 75° 5' W., while the eastern margin is approximately defined by the extensive salt marshes along the bays and channels shut off from the sea by barrier beaches (Colored map, Plate 1). In a northeast-southwest direction the region is about 1 2 1 kilometers (75 miles) long and from west to east in its broadest portion approximately 57 kilometers (35 miles). The coastal plain of this region is rolling with ridges, valleys and slopes, but notwithstanding that fact, when one views the country from an ele- vation he discovers its essentially flat character. So level is it in places that the greater height of the cedar trees in the swamps impresses the uninitiated as a ridge or hill covered with coniferous trees. The coun- try is still in many places an unbroken wilderness. The drainage area of the Wading River is one of the wildest, most desolate portions of the eastern United States, and the writer has traveled a distance of 16 kilometers (10 miles) without seeing a house or a human being. Throughout the soil is a silicious sand underlaid with red colored gravel deposits. The fall of the eastward-flowing streams is quite uniform. Above the head of the tide the average fall of the Manasquan is 1.6 meters (53^ feet) per mile; Toms River, 1.9 meters {6j4 feet) per mile; Cedar Creek falls 2.1 meters (7 feet) for 11.2 kilometers (7 miles); Mullica River, 1.5 meters (5 feet) for 26.7 kilometers (16 miles), and Great Egg Harbor River, i.$ meters (5 feet) for 40.2 kilometers (25 miles).* The water of these streams is a pale coff'ee color and is known as cedar water. Its color is due to organic matter leached out of the soil and the peaty deposits of the swamps. Origin of the Designation "Pine-Barrens" With the general physiography and the general distribution of the vege- tation of coastal New Jersey in mind where the forest of pine trees oc- *Vermeule, C. Clarkson: Physical Description of New Jersey. Final Report i, Geological Survey of New Jersey, 1888: 175. VEGETATION OF THE NEW JERSEY PINE-BARRENS 9 cupies the central portion of the state and the deciduous vegetation the richer alluvial soils of the streams which flow into the Delaware River, as well as the richer interstream areas, we have an explanation of the designation "barrens" applied to the region of the pitch-pine forest. When the country was discovered first, before clearings and roads were made in the primeval forest, the early settlements were made along the larger streams, notably the Delaware, where the soil was rich and productive. After all the available land along the larger rivers had been patented, the early settler utilized the unsettled wilderness to the east- ward. As the population became denser it encroached more and more upon the region covered with the forest of pitch-pines, and as the soils of this region were thinner and less productive, the region was looked upon as barren. Soon the name "barrens" was used by the early im- migrants, and this name was coupled with pine, giving the well-known epithet of "pine-barrens." The word barren is used elsewhere as a de- scriptive geographic term. There are serpentine barrens in southeast Pennsylvania, where the unproductive outcrops of serpentine rock are distinguished by the presence of plants peculiar to that type of soil. Barren Hill, near Valley Forge, Pennsylvania, was occupied by the Con- tinental soldiers on May i8, 1778, in order to oppose the advance of the British. The barren grounds of Canada represent the flat arctic coun- try known as the tundra. All of these names are significant of the relative poorness of the land. Similar areas of pineland exist south of Virginia along the southern Atlantic and Gulf coastal plains and these have been named pine-barrens. But the plant geographer soon distin- guishes several kinds of pine-barrens, which may be classed as follows: Pitch-Pine (Pinus rigida). Cape Cod, Long Island and New Jersey. Loblolly-Pine (Pinus taeda), Virginia to Florida and Texas. Long-leaf Pine (Pinus palustris), southeastern Vir- ginia to Florida and Texas. Slash-Pine (Pinus caribaea), Florida to South Caro- lina, westward to Mississippi, Bahamas, and Cuba. Yellow Pine (Pinus echinata), Arkansas and north- . eastern Texas. Geographic Place Names Much has been made of the study of place names as indicative of the early settlement of countries by men of different nationalities and races. * Forthe characterization of the different kinds of pine-barrens consult Harshberger, J. W.: A Phytogeographic Survey of North America, 191 1, vol. xiii, Die Vegetation der Erde. Pine-Barrens* 10 VEGETATION OF THE NEW JERSEY PINE-BARRENS When all trace of these peoples has been lost in a neighborhood, the names given to rivers, lakes, hills, and towns still attest to the former occupancy of the country by the people who bestowed the name as a geographic landmark. In this country considerable study has been made of the Indian names of places,* but the significance of the names as given by the early settlers has been touched barely by American philolo- gists. Thus in England the name of the City of Chester is indicative of the location of a Roman camp, castra. In Pennsylvania, the location of the early Welsh towns is shown by such place names as Bryn Mawr, Cynwyd, Gwynedd, Radnor, Ithan (a creek), and the like. Selections of the characteristic place names in the pine-barren region of New Jersey which are indicative of some local peculiarity and with special reference to the flora are here given. The white-cedar tree (Chamaecyparis thyoides) has been used in designating a considerable number of geographic features of the region, as: Cedar Bridge, Cedar Brook, Cedar Grove, Cedar Lake, and Cedar Run. Next to the cedar the white-oak has been used, as White Oak Bottom, White Oak Branch. Then a few well-known trees and shrubs have been chosen for place names, as: Chinquapin Branch, Gum Spring, Huckleberry Hill, Mag- nolia, Maple Root Branch, Pine Creek and Red-Oak Grove, while general impressions of the vegetation are expressed in such names as: Grassy Pond, Great Swamp, Green Branch, Wood Swamp. Names which have been applied during logging operations are Log Swamp Branch, Pole Branch, Rail Branch, Slab Causeway Branch and Tarkiln Branch. Indian names, although relatively little used, are found to some extent on the maps of the region. Such are Indian Jack, Mana- hawkin, Manopaqua Branch, Manumuskin, Mecheseatauxin Creek, Muskingum Brook, Nesochoque Branch, Papoose Branch, Shamong (Chemung = big horn), Tulpehocken Creek, Tuckahoe, Westecunk River. The following place names commemorate well-known animals. Those of exotic introduction are noted by italics: Bass River, Bears Head, Bear Swamp, Beaver Branch, Beaverville, Bull Branch, Bulltown, Chicken Bone, Fox Chase, Goose Pond, Mare Run, Panther Branch, Turkey Foot, Wild Cat and Wolf Run. Names of places that express physiographic conditions are: Deep Run, Forked River, Great Swamp, Goodwater Run, Sandy Ridge, Wading River. A number of geographic features are indicated by well- known articles of human use, as: Apple Pie Hill, Bread and Cheese Run, Breeches Branch, Cabin Branch, Calico, Featherbed Brook, One Hun- * Beauchamp, William M.: Aboriginal Place Names of New York. New Yori< State Museum. Bull. io8, 1907. VEGETATION OF THE NEW JERSEY PINE-BARRENS II dred Dollar Bridge, Tub Mill, and some have a personal flavor, viz.. Comical Corner, Double Trouble, Friendship Bog, Good Luck, Hospi- tality Branch, Jake Branch, Jemima Mountain, Mary Ann Furnace, Mount Misery, Pleasant Mills, Stop-the-Jade Run. The following few places have been designated by terms expressive of distance or dimen- sions: Five-Acre Pond, Four-Mile Run, Old Halfway, Ten-Mile Hol- low. The origin of such names as Aserdaten, Atco, Atsion, Batsto, is unknown to the writer, but in the miscellaneous assortment the follow- ing are noteworthy: Burnt Bridge Branch, Head of Snag, Hurricane Brook, Landing Creek, Long Cripple, Money Island, New Freedom, Oriental, Pole Bridge Branch, Stormy Hill and Tabernacle, Some of the geographic names in the pine-barrens are names of persons compounded with forge, mill, town, ville, as: Bennett Mill, Buddtown, Hance Bridge, Hesstown, Monroe Forge, Smithville and Webb Mill. How much more applicable and appropriate the names mentioned above are than the following examples of exotic ones that have been transplanted to the pine-barren region: Chatsworth, Chesilhurst, Lake- hurst, Penbryn. Railroad officials, county officers and the post office authorities should be very careful not to replace an old, well-established and expressive name which has a local flavor for a "high-falutin" term of no local significance.* * GiFFORD, John: Forests of New Jersey, 241, footnote. CHAPTER III INFLUENCE OF GEOGRAPHY AND VEGETATION ON THE PEOPLES AND THE INDUSTRIES OF THE REGION General Considerations The geography of any country modifies very profoundly the settlement, degree of civilization and industries of that country. Nowhere is that principle illustrated more clearly than in the pine-barren region of New Jersey. Until the opening of the first railroad through the region, travel was by horseback, wagon, or coach. The roads were cut through the wilderness and were few and far between. The inhabitants of the pine forest were isolated from their neighbors and by long isolation they have developed racial peculiarities of dress, speech and thought which are today characteristic. At present from an elevated hill we see an unbroken extent of dark-green pine forest, as far as the limit of vision, stretching away in long gentle swells. Much of the region is an unbroken wilderness save for the damage that has been wrought by forest fires and abandoned clearings scattered through the forest. Writing in 1885, Vermeule* says: "From Manchester southward to the Mullica River is one of the wildest, most desolate portions of the State. If we except the clearings on the shore road and along the marl border, not more than 2 per cent of the area is under cultivation. Here and there are narrow roads, barely wide enough for a single vehicle to pass clear of the trees, which thread their lonely way from clearing to clearing. They are relics of a time when the manufacture of iron from bog ore found in the swamps was an important industry of the region. Here and there one comes upon abandoned forge sites, or still more suggestive, aban- doned villages, the relics of unsuccessful glass manufacture in the wil- derness. An indescribable silence prevails. The soughing of the wind through the pines oppresses, while the crowing of a cock or the barking of a dog indicates the approach to a clearing and human habitation." If the above quotation describes the condition of the pine-barrens in 1885, what must have been the character of this wilderness when the country was settled? So inaccessible and difficult was the travel across * Vermeule, C. Clarkson: Final Report of the State Geologist, 1885, vol. 1: 177. 12 VEGETATION OF THE NEW JERSEY PINE-BARRENS 1 3 the continuous pine forest and so little was to be gained by settlement that the cultivable land was utilized very slowly by the early white settlers. Even the Indians of South or West Jersey rarely penetrated these pine- barrens except for hunting or fishing, for at the arrival of the white men we find the tribes of the Lenni-Lenape stock inhabiting the Dela- ware River Valley along both the east and the west banks. The most southern tribes, in what are now Salem and Cumberland counties, New Jersey, belonged to the Turkey Totem, while those opposite Philadelphia and extending to Trenton belonged to the Tortoise Totem. No tribes lived on the Atlantic coast of New Jersey, according to early authorities, and the shell heaps along the shore were formed by the Indians who now and then crossed the pine-barrens to enjoy the sea-fishing and the gather- ing of the shell-fish. The topography of the country influenced the location of the early settlements which were made along the Atlantic coast within easy access of the ocean by the coastal rivers and bays by light draft boats, and along the Delaware River, all points of which were easily accessible by vessels of greatest draft at that early day of sailing vessels. The principal settlements made in West Jersey before the country became a royal colony in 1703, and while in possession of Hol- land and Sweden (1609-1664); of Berkeley and Carteret Proprietary (1664-1673); of Province (east and west division, 1673-1703) were, in the order of the date of their foundation: Shrewsbury, 1664; Barnegat, 1668; Nevesink, 1670; Burlington, 1674; Manasquan, 1684; Mount Holly, 1685; Cape May Town (Portsmouth, Town Bank), 1686; Ran- cocas, 1686; Egg Harbor (Tuckerton), 1690; Bridgeton (Cohansey Bridge), 1690; Somerset Plantation (Somers Point), 1695; Fairfield, 1698; Cohansey (Caesarea River), 1698; Haddonfield, 1701. This vast, irregular-shaped tract of land, the " Pines," stood as a barrier between the early settlements on the Delaware and those along the sea-coast, where fishing and whaling were important industries.* Roads designed to connect these early settlements were early con- structed across the pine-barren wilderness, where abounded red deer, bears, wolves, panthers, wild-cats, foxes, rabbits, opossums, pole-cats, hedgehogs, wild turkeys, pheasants, grouse and quails. Wolves were so abundant that a bounty of ten shillings was offered in 1682 for every head of that animal killed in West Jersey. f Earlier than these roads several Indian trails^ led through the pine- * Harshberger, J. W.: The Hardy Fishermen of New Jersey. Old Penn, xi: 773, March 25, 1913, with 7 figures. t Lee, Francis Bazley: New Jersey as a Colony and as a State, i: 286. t Proceedings of the Surveyors' Association of West Jersey: 408. 14 VEGETATION OF THE NEW JERSEY PINE-BARRENS barrens. One started from Somers Point and extended along the east side of Great Egg Harbor River, so as to strike north of the heads of the several branches of Babcock Creek; and over the lowlands of that branch as it approached the tributaries of Little Egg Harbor, called the " Locks, " by Blue Anchor and crossing the head of the Great Egg Harbor River at Long-a-coming (Berlin), passed a short distance south of Had- donfield, striking the Delaware at Coopers Point. The second Indian trail came from the mouth of Little Egg Harbor River in a westerly direction and joined the first-mentioned trail near the head of Landing Creek, one of the tributaries of the last-named stream (colored map). The third began near MuUica's plantation near Batsto and went in a westerly direction between the streams to the first-named trail at the old Beebe place, about 1.6 kilometers (i mile) south of Winslow. The fourth trail, later called the Old Cape Road, started in Cape May County. It crossed the head of Tuckahoe River in a northerly direc- tion and to the west of the branches of the Great Egg Harbor River to the upper waters of Hospitality Creek at Cole's Mills, thence to Innskeep Ford, and joined the first-named trail at Blue Anchor, which was a cen- tral point. These Indian trails were used by the white settlers between the Delaware River and the coast until they were converted into roads suitable for wagon and coach travel, while other highways were sur- veyed and built through the pine-barrens, which in the eyes of the pro- prietors of West Jersey were of comparatively little value. By 1700 the pine-barrens had been crossed by surveyors and much of the timber had been purchased from the proprietors.* The roads which ran through the pine forest so as to reach the timber grants, as well as the settlements on the east coast, were generally sandy, and during the dry season the tires of the wheels pulled heavily through the loose surface. One of the most important of the later roads con- nected with the shore road at Barnegat. It ran northwestward through what are now Cedar Grove, Woodmansie, Ong's Hat, South Pemberton, Birmingham, Mount Holly, and reached the Delaware at Burlington. These highways opened up the country to the earlier English settlers whose names, as derived from ancient land grants and from their de- scendants, with whom the writer has come in contact, were Bartlett, Brown, Crane, Cranmer, Gaskill, Haven, Jennings, Moore, Seaman, Shaw, Southwick, Truax, and Wainwright, while at Barnegat we find the family names Arnold, Birdsall, Bird, Collins, Inman, Loper and Mott.f * Lee, Francis Bazley: New Jersey as a Colony and as a State, i: 283. t Blackman, Leah: History of Little Egg Harbor Township, Burlington County, from its First Settlement to the Present Time, 1868. VEGETATION OF THE NEW JERSEY PINE-BARRENS I 5 Considerable space has been devoted to this historic sketch of the early settlement, because we have reason to believe that that settlement was influenced by the character of the country and in turn profoundly modi- fied the original boundaries of the pine-barren region. As the land along the shore road was converted into farms at a very early date, perhaps prior to March, 171 3-14, when, owing to the excessive destruction of the forest, an act was passed prohibiting the common practices of stealing timber, cedar, pine staves and poles and of boring for the extraction of turpentine, the natural succession of the forest growth must have been greatly altered. And these changes occurred not only in the coastal strip of country back of the salt marshes, but in Cape May County as well. Floral Districts Stone,* in his book previously cited, recognizes five distinct floral dis- tricts in southern New Jersey: (i) The West Jersey, or Middle District, which covers, according to his colored map, not only the Delaware Valley region south of Trenton, but also all the country below the fall- line and north of the pine-barrens which terminate at Long Branchf; (2) the Pine-Barrens; (3) the Coastal Strip; (4) the Cape May District southoftheGreat Cedar Swamp; (5) the Maritime District. Stone'sclas- sification of districts is somewhat empiric. Having considered the his- tory of the settlement of South Jersey, we are in a position to understand that before the year 1725 serious inroads had been made upon the forests, especially along the coast and on the peninsula of Cape May, where a number of flourishing towns had been started. These towns were largely inhabited by whalers and fishermen, who found ready access to the sea from the mainland. Farms were cleared in the pine forests, which probably extended down to the salt marshes. As the whaling and other industries of the sea declined in the Cape May peninsula and along the New Jersey sea-coast, many of these farms were abandoned and the open fields were invaded by deciduous trees, quite distinct from the pine trees which originally covered the country. Such settlement undoubt- edly disturbed the original conditions, for the writer, in a former publi- cation,! has referred to the influence of cattle on the forest at Wildwood. Cattle for many years ran wild on this island, which was granted by Charles II of England to his brother James, Duke of York, on March 12, * Stone, Witmer: The Plants of Southern New Jersey, 57. t More correctly. Deal Lake. t Harshberger, J. W.: An Ecological Study of the New Jersey Strand Flora. Pro- ceedings Academy of Natural Sciences, 1900: 630. l6 VEGETATION OF THE NEW JERSEY PINE-BARRENS 1664. These cattle must have roamed for two hundred years over the island until they were shot only a short time ago. Many of the peculiar growths at Wildwood (now completely destroyed) were due undoubtedly to the trampling down of the young growth and the browsing of animals upon the tender buds and shoots, so that this irregular method of prun- ing and training caused the trees to assume curious forms and produced natural graft unions. The Cape May District and the Coastal Strip which Stone assumes are natural districts are, if our observations are correct, of artificial production. They exist today as well-recognized phytogeographic areas, but judging from the fact that in several places today the pine forest touches the salt marshes, they were covered by a pine forest integrally connected with the pine-barrens inland. If this surmise is correct, then theoretically we can reduce Stone's floral districts to four by excluding the Coastal Strip. If all of the peculiar plants known to exist in Cape May County, as well as the components of the deciduous forest, have been introduced with that forest type since the settlement of the country, then the Cape May District as distinct from the Pine-Barren District ceases to exist, reduc- ing Stone's districts to three. The Cape May vegetation gives a strong impression of a mixed character, which would follow the occupancy of abandoned fields by pioneer vegetation of the deciduous type. As a result of this analysis the floral districts of South Jersey are three: (i) The Middle District; (2) the Pine-Barrens; (3) the Maritime District, so that the red of Stone's map would on theoretic grounds be extended on the east to the green color of the salt marshes, and on the south to the tip of the Cape May peninsula. Considering the encroachment of cul- tivated areas upon the pine-barrens, the region in all probability ex- tended farther west toward the Delaware River. The influence of the geographic environment on the development of the character and civilization of the inhabitants of a region is deter- mined best by a study of their outlook on nature and by a consideration of their industries. We are concerned in this monograph with these problems, as far as the vegetation of the region has developed the powers of observation and habits of the people, or has influenced daily life and the development of the industries for which the region is noted. Local Plant Names Local names survive longest, as a rule, in the process of speech evolu- tion, and bear evidence of the cultural stage and inventive genius of a people in introducing new terms for new phases of nature and in describ- ing new animals, plants, and geographic places (see ante). The follow- VEGETATION OF THE NEW JERSEY PINE-BARRENS 1 7 ing is of interest:* The fishermen along Little Egg Harbor River call the eel-grass (Zostera marina) tiresome-weed, for, as they pull against the current in going to and from the ocean, the grass-wrack entangles their oars and retards the movement of their boats. The people in some districts in the pine-barrens call the orchid, Cypripedium acaule, "whippoorwill-shoe," which is better than that of lady's-slipper. The common pitcher-plant, Sarracenia purpurea (side-saddle-flower, huntsman's-cup), is called locally in Cape May County "dumb-watches." The flower grows from between the rosette of pitchered leaves, and after a time the incurved petals fall, exposing the top of the broad, ex- panded stigma. The five sepals remain and resemble, to the imagi- nation of the New Jerseyman, a watch-case, while the convex surface of the stigma is likened to the face of a watch. This vegetable time- piece, with no hands to point the hours of the day, without the constant tick, tick, tick, is dumb. The whole plant is denominated very significantly and ingeniously "dumb-watch," or simply "watch." Other examples of this outlook upon nature might be cited, but sufficient has been given to indicate that a person interested in folklore would find an interesting field for investigation among the simple men and women of the pine-barrens. Industries of the PiNE-BARRENsf With the exception of the mining of bog-iron ore associated with the construction of ore furnaces and the utilization of the finer clay deposits and glass sands, the industries of the region from the earliest times have been dependent on the products of the forest and of the bogs so char- acteristic of the country. Lumbering. — The saw-mills driven by water power heralded the advent of permanent settlement. The first saw-mills were erected in the period 1700-1725 (Figs. 3 and 4). An act was passed in 1743 laying duties upon logs, timber, planks, vessel supplies, staves and headings, except firewood, exported to any of his Majesty's colonies upon the continent of America. The pitch-pine, Pinus rigida, is the tree which gives character to the pine-barrens of southern New Jersey. It grew originally from 12 to 32 meters (40 to 75 feet) tall in the primeval forest, but at the present day, owing to frequent forest fires, it is rare to find trees as tall as 1 5 meters * Harshberger, J. W'.: Local Plant Names in New Jersey. Garden and Forest, v: 395. 1892. t See a suggestive paper along entirely different lines, W'hitbeck, R. H.: Geographical Influences in the Development of New Jersey. Journal of Geography vi: 177-182, 1907- 08. VEGETATION OF THE NEW JERSEY PINE-BARRENS (50 feet). The tree as usually found ranges in height from 6 to 12 meters (20 to 40 feet). The trunk can be manufactured into coarse lumber, and in the smaller sizes is made into charcoal or cut into firewood. It has never been a favorite tree, owing to the poor quality of the timber, but yet it supplied a want in the early days before the pineries of the southern states were exploited, especially as it was possible to cut large trees that yielded straight and fair-sized boards, joists, and scantling for the construction of log houses (Fig. 5). One or two hundred years ago, when the best pitch-pine lumber could be obtained, it was employed principally for the sills and beams of buildings. Structurally the wood is soft, light, coarse-grained, and durable, and although little used for construction at present, the larger sizes can be used for rail- road ties. The ease with which the pitch-pine can be raised from seed, and its rapid growth on sterile soils, make it of especial value in silvicultural operations, where it is desired to cover barren areas with a forest cover, as in the Island of Nantucket and the dune sands on Cape Cod. The early saw-mills (Figs. 3 and 4) were driven by water power derived from the streams which were dammed at favor- able localities accessible to the forest, where the trees were cut by the axe. As compared with modern logging operations in the west and south, the methods of cut- ting and handling the logs in the forest over one hundred years ago were of the simplest description. Horse and ox teams sufficed to trans- port the logs to the saw-mill, and the largest sizes were handled easily by one or two men. As the country is level and flat, no insuperable dif- ficulties were presented in getting the logs out of the woods, especially if the lumbering was done in the winter, when the frozen ground was cov- ered with snow. Sleds drawn by ox teams could easily remove the logs to the saw-mills driven by an overshot or an undershot waterwheel. Some of these old saw-mills are still used and are of great interest to the historian of the lumber industry in America. The early builders of mills in New Jersey discovered that pitch-pine was well suited for the Figure 3 Old colonial saw-mill erected in 1750 and in continuous use since on Darby Creek, near Manoa, Pa., taken May 2, 1914. Similar saw-mills were in operation in New Jersey and were scattered through the pine-barrens, being driven by water power. VEGETATION OF THE NEW JERSEY PINE-BARRENS 19 FlGUKU 4 Interior of old colonial saw-mill erected in 1750 on Darby Creek, Pa. The band saw with up-and-down movement is driven by wheels and other parts constructed of wood. 1 he log-carrier with partly sawn log is shown in the foreground. May 2,1914. buckets and spokes of the waterwheels, which worked partially under water, because as a wood it re- sisted decay. The heartwood was used for boat building and for ship pumps.- In Pennsyl- vania pitch-pine logs are used as mine props, wharf and bridge piles. Pitch-pine floors* wear well, look well, and last a long time. A pitch-pine floor in Pike County, Pennsylvania, laid with boards 0.6 meter (2 feet) wide and 3.8 centimeters {i}4 inches) thick, did service one hundred and sixty years and was still in such a good condition that the boards were used in flooring a new house. t The wood has been used suc- cessfully for door and window frames, ceiling and other interior finish, and although knots are fre- quently abundant, yet, if prop- erly finished, they give a decor- ative appearance to the wood. 1 1 is used to advantage for wood- pulp, excelsior, barrel headings, crates, nail- kegs, beds for wagons, and the inside boards of cupboards, ice-chests, cab- inets, tables, and desks. One of the largest uses for pitch- pine is in the manufacture of boxes and crates. t In 1909 the box-makers of Massachusetts used 600,000 feet, and Mary- land, 61 5,000 feet, of pitch-pine. As cordwood for fuel it has sup- FlGURE 5 Old colonial log school house at Speedwell, Woodland Township, Burlington County, showing style of old pine-barren architecture and construction. (Courtesy of Herbert N. Morse.) * Hall, William L., and Maxwell, Hu. : Uses of the Commercial Woods of the United States. H. Pines, U.S. Forest Service Bulletin, 99: 34, 191 1. t Defebaugh, J. E.: History of the Lumber Industry in America, 11: 563. t Hall and Maxwell; Loc. cit. 20 VEGETATION OF THE NEW JERSEY PINE-BARRENS plied brick-kilns, potteries, bakeries, steam engines, and houses for many years. Fat pine-knots, which were obtained from old rotten logs by knocking away the decayed wood, or which were found in the forest in an excellent state of preservation after the softer, less resinous wood had disappeared, were used as a substitute for candles and lamps in the early days. These knots, which were gathered as fuel for winter's use, were split into thin pieces for illuminating purposes. These were bound into bundles with hickory, or yellow-birch, withes and were used as torches in night travel through the woods. "The dwelling-rooms of the Swedish settlers were lighted partly with tallow-candles and partly with so-called perstickor, or spingstickor. These were splint-sticks of pine used very extensively in this country and in Finland and Sweden, being mentioned in the Kalevala. They were thin and flat and were made by splitting the pine knots into proper size. They were fastened into a stick-holder of iron with a clasp for holding the pine splint and a sharpened end to stick into the crevices between the logs or into the wall. The other type was an upright shaft of wood, a yard or more in height, which could be moved from place to place. The splint was attached with the free end pointing downward, which was lighted. As rapidly as these torches were burned down they were replaced by fresh ones. Two or three burning sticks lighted a room for ordinary purposes, but they produced much black smoke."* The spearing of fish at night was facilitated by the use of pitch-pine torches. Charcoal was manufactured also from the inferior grades of pine logs and it was used in blacksmith shops and iron forges quite extensively. Lamp-black was made from pitch-pine. The cedar swamps yielded and still furnish a fine lumber obtained from the white-cedar tree (Fig. i). The durable qualities of white- cedar, especially in contact with the soil, were early appreciated. Al- though the wood is soft and rather weak, yet on account of its non- resinous character and its straight grain, it is an ideal material for boat- building, for cooperage (vats, tanks, churns, piggins, firkins, tubs) and various kinds of woodenware. Gottlieb Mittelberger, a German wagon- builder, considered the white-cedar as one of the finest materials for the construction of church organ pipes. Shingles made of white-cedar last many years. There is a specimen of a split shingle in the wood collec- tion at the University of Pennsylvania, taken by the writer from the side of a house on Long Island built probably for at least two hundred years. The wood has been used satisfactorily for interior finish (floors, doors, * Johnson, Amandus: The Swedish Settlements of the Delaware, 1638-1664, vol. i: 351; Retzius: Finland, p. 73 ff. VEGETATION OF THE NEW JERSEY PINE-BARRENS 21 joists, frames, rafters), fence posts, telegraph poles, and railroad ties. Coincident with the lumbering operations in the pine forest, the cedar swamps were culled of their largest and best trees. Even today, after two hundred years of such exploitation, fair-sized cedar logs are removed from the forest. According to Hall and Maxwell,* the cutting of south- ern white-cedar began probably three hundred years ago and was in full blast in New Jersey two centuries ago. John Lawson, nearly two hun- dred years since, mentioned its use in the Carolinas for "yards, top- mast booms, and bowsprits for boats, and shingles and pails." The drain on the swamp forests for white-cedar lumber was so great that Benjamin Franklin published an essay in Poor Richard's Almanack (1749) in which he advocated forestry methods, especially the planting of red-cedar to supply the country when the white-cedar and other woods should fail. Peter Kalm foretold the inadequacy of the white- cedar swamps to meet the demands for its excellent timber. But the supply is still far from exhausted. The indestructibility of the cedar wood is shown by the almost perfect state of preservation of logs that have been buried naturally in the cedar bogs. The peaty soil is soft and spongy, and in some of the older bogs the swamp bottom for many feet is a tangled mass of fallen logs between which is found peaty material of a more or less decayed character, con- sisting of the remains of leaves, twigs and moss. In some bogs these buried logs have been raised by mining operations, the position of the logs having been determined by probing with an iron rod. The location of the logs by the iron rod is followed by the digging of a trench so that the log can be floated to the surface. Many valuable pieces of timber have been secured by these mining operations. The water percolating through the cedar swamps emerges with a light brown or tea color, and has a faintly pungent cedar taste. As this water has antiseptic properties, it was long ago valued by sea captains, who filled their casks with cedar-water before going on a long voyage. Some one had discovered that cedar-water kept sweet and potable longer than ordinary river-water. Turpentine Industry. — The oleoresin which exudes from pine trees is known as crude turpentine, and it is obtained by incisions made in the tree, which is said to be boxed. The early box system of gathering tur- pentine was imperfect and wasteful. Boxes about lo centimeters (4 inches) deep and 20 centimeters (8 inches) wide, according to the size of the tree, were cut in the living trees about 30 centimeters (i foot) from * Hall, William L., and Maxwell, Hu.: Uses of Commercial Woods of the United States. I. U. S. Forest Service, Bulletin 95: 13,1911. 22 VEGETATION OF THE NEW JERSEY PINE-BARRENS the ground. The trunk was scarified by slanting cuts above the box by an axe especially constructed for the purpose (Fig. 6). The chipping began in March and continued well into the summer. The oleoresin, which exuded from the stem, followed the converging, slanting cuts into the box. A laborer could cut one hundred boxes in a day (Fig. 7). The gathering, or the dipping, of the crude turpentine began three or four weeks after the chipping. The first gathering was known as the virgin dip, and subsequent collections were made through the season. At the end of the flow there was an accumulation of dried turpentine called " scrape," because it was scraped off into receptacles carried through the forest for the pur- pose. The crude oleoresin was then taken to the still (Fig. 8). The old method of cutting deep boxes (Fig. 7) proved very des- tructive to the tree and reduced the term of operation to three or four years. Recently a more economic method of collecting the crude oleoresin has been adopted through the experi- ments conducted in the forests of the south by the U. S. Forest Service.* It is known as the cup-and-gutter method, where small galvanized iron gutters are attached in a slanting direc- tion at the base of the scarified surfaces and the exudation runs down into earthen cups attached by a nail at a point where the gutters converge. The amount of crude resin collected is greater than by the old system. It is cleaner and the crude turpentine is more expeditiously handled. The old method, however, was the one used in New Jersey when Gabriel Thomasf Figure 6 Primitive method of boxing pine trees to obtain crude turpentine as practised about 1700 in New Jersey. (Photograph (No. 40,162) taken near Ocilia, Georgia. Cour- tesy of U. S. Forest Service.) * Consult A New Method of Turpentine Orcharding. U. S. Bureau of Forestry, Cir- cular No. 24; Herty, Charles H.: A New Method of Turpentine Orcharding. U. S. Bureau of Forestry Bulletin 40, 1903; Herty, Charles H.: Practical Results of the Cup and Gutter System of Turpentining. U. S. Bureau of Forestry Circular No. 34; Davis, Charles: Making the most from Pine Orchards. The Country Gentleman, Aug. 3 i, 1912. t Thomas, Gabriel: An Historical and Geographical Account of the Province and County of Pennsylvania and West Jersey in America, 32. VEGETATION OF THE NEW JERSEY PINE-BARRENS 23 wrote of Robert Stiles (d. 1713), who lived on a farm on Pensauken Creek in Chester Township, Burlington County, just over the line from Gloucester, that " the Trade in Gloucester County consists chiefly in Pitch, Tar and Rosin; the latter of which is made by Robert Styles, an excel- lent Artist in that sort of Work, for he delivers it as clear as any Gum Arabick." Crude turpentine* is distilled now in copper stills. Formerly the use of iron stills was universal and water was used in the distil- lation (Fig. 8). A fifteen-barrel still (barrel weighed 127 kilos — ■ 280 pounds) was charged early in the morning. Gentle heat was applied first until the mass was in a liquid state, when a coarse wire skimmer was used to remove the chips, the bark, the leaves, and such other foreign substances as rose to the sur- face. Meanwhile the tempera- ture rose until 316° F. was reached. All the accidental water (that contained in the crude oleoresin as it came from the forest) having been distilled off, a small stream of water was directed into the still, so that the heat did not rise above 316° F., the boiling-point of oil of turpentine. The oil of turpen- tine and water distilled over, and the mixture was caught in a wooden tub. The distiller tested the quality of the flow from time to time and the distillation was continued until the proportion of water coming over was nine of water to one of turpentine. At this stage the heat was withdrawn, the cap of the still was taken off, and the hot rosin was drawn off by a valvular cock at the side of the still near the bottom. The rosin was Figure 7 Primitive method of boxing pine trees to obtain crude turpentine, as practised about 1700 in New Jersey. (Photograph (No. 40,168) taken near Ocilla, Georgia. (Cour- tesy of U. S. Forest Service.) * Consult Hawley, L. F.: Wood Turpentines. U. S. Forest Service Bulletin 105, Washington, 1913. 24 VEGETATION OF THE NEW JERSEY PINE-BARRENS passed through a strainer before it reached the barrel to rid it of foreign substances. In the early days, when turpentine was made in New Jersey, besides the turpentine and rosin, which were the main products, pitch was also obtained. Rosin when subjected to dry distillation yields a number of products: resin-oil (85 per cent.); resin-spirit; water; a powerful anaes- thetic gas with a residue of pitch* amounting to i>2 cwt. from a ton of rosin. Shoemakers' wax was made by melting together the best pitch and tallow in a vessel over a fire. The resulting mass was rolled into balls and was used to wax the linen thread used by cobblers — the so-called "wax ends" being formed. Tar (Pix liquida) was used by country people in the early his- tory of this country as axle grease, and the wagon without the tar bucket and its tar paddle swinging from the rear axle was an oddity. Tar was manufac- tured by the backwoodsmen of New Jersey by using stumps, roots, and other waste mater- ials of the pitch-pine lumber- men. The important features in tardistillation was the pile of wood which had to be slowly and imperfectly burned from the top down- ward, so that the tar would be driven successively out of the wood next underneath the burning point, running down and out at the bottom, where it was collected. The stacks of pine refuse consisted of circular mounds of earth about 6 meters (20 feet) in diameter, hard-packed and clay-coated, surrounded by a ditch used to collect the tar as it was formed. As the wood was piled upon the mound, the diameter of the pile gradually increased to about 8 meters (25 feet) at the top and some 3 to 4 meters (10 to 12 feet) high. The top was then covered with pine needles, and the sides of the heap were protected against a free current of air and the top ignited. The tar was barreled as fast as it ran away. Such a stock burned for a week or ten days and yielded 100 barrels or * For properties of rosin-pitch consult Hopkins, Albert A.: The Scientific American Cyclopedia of Receipts, Notes and Queries, twenty-second edition, 1903: 606. Figure 8 Primitive form of turpentine still as used in New Jersey about 1700. (Photograph fur- nished by U. S. Forest Service (No. 15,018), taken at Clinton, Sampson County, North Carolina, by Worth.) VEGETATION OF THE NEW JERSEY PINE-BARRENS 25 more of tar, which was a thick, viscid, semifluid mass, heavier than water, and of a deep, blackish brown color, having an unpleasant empyreumatic odor, bitter taste and of a complex chemic composition. Another slightly different method employed in the manufacture of tar was used during the Civil War at Tuckerton, New Jersey, where it was a profitable industry. A large basin was constructed from which a small trough, or channel, led into another receptacle, and the whole was carefully cemented with a composition of sand, loam, and clay. The pine knots were piled in the large basin in a cone-shaped stack and cov- ered with soil, sod, etc., similar to the preparation of a charcoal pit, and then lighted at the top. The tar oozing out passed into the bottom of the basin, thence through the channel into the receiver, from which it was collected.* Cranberry Culture. — An important industry of southern New Jersey which illustrates what the natives of the region have done in the domestication and cultivation of a native wild plant is the cultivation of the cranberry, Vaccinium macro- carpon,t for the fruit of which a large demand has developed in the United States, for the total product in 1903 was 344,986 hectoliters (977,516 bushels). The soils for the successful culture of the cranberry must be of a peaty character, with the water level within a few inches of the surface of the soil. The supply of water should be sufficient and the plantation so provided with dykes as to allow of flooding the area with water to the depth of 46 centimeters (18 inches) to 61 centimeters (2 feet) from November to May, in localities where it is necessary to protect the plant from insects and from late spring frosts. In the preparation of the bogs, many of which exist in southern New Jersey (Fig. 9), all bushes and trees must be removed, together with the top layer of soil to the depth of from five to ten centi- meters (2 to 4 inches). After turfing, the surface must be graded, dams Figure 9 Cranberry bog and drainage ditch at Browns Mills. (Photograph by Henry Troth.) * HiLLMAN, Sarah C: The Pine Lands in their Flora and Fauna. Proceedings Dela- ware County Institute of Sciences, iii: 1 14-123, April, 1908. t Harshberger, J. W'.: Peat Bogs and Peat. Old Penn Weekly, April 10, 1909, page 8. 26 VEGETATION OF THE NEW JERSEY PINE-BARRENS and sluice gates must be constructed, and ditches dug to facilitate drain- age. The surface must be sanded to a depth of 8 to lo centimeters (3 to 4 inches), and the young plants must be set out as cuttings by dibbling the soil in rows about the first of June. The only cultivation necessary is to keep down weeds and grasses, which are not troublesome where the bogs are flooded periodically. Harvesting the fruit, formerly done by hand, is now performed in some places by a cranberry scoop-rake, the sorting being done in sorting racks in which the berries are separated into the different marketable sizes. The sorting sheds are low, rambling affairs, located in temporary villages in the higher pine-land, or larger, more substantial buildings (Fig. 10). As the diseases of the cranberry have increased of late, agents* of the United States Department of Agriculture have investigated them with a view to overcom- ing the fungi and insects which attack the plant. Much good work has been accomplished. Uses of Sphagnum Moss. — There are many bogs in southern New Jersey where bog moss, or sphagnum, is gathered for use in packing plants for shipment by wagon and railroad, and is baled for shipment to gardeners and nurserymen. It is used in the cultivation of orchids, ferns and pitcher-plants, which re- quire a constant supply of water about their roots for their best development. Sphagnum is used in the operations of grafting and propagation of such plants as the rubber plant (Ficus elastica), which will produce new roots whenever a fistful of bog moss is tied around the stem, which, after the adventitious roots have appeared, is severed below the ball of moss. Peat. — The sphagnum bogs of New Jersey yielded peat, but of an inferior grade. Owing to the structure of the moss, the bog is satu- rated with water. The upper extremities of the moss stems continue their growth from year to year, while the lower portions die away and are converted gradually into peat by the pressure of the wet moss above Figure 10 The Birches. Cranberry sorting and storage house, with wagon loaded with cranberry bar- rels. Note martin houses on poles. Taber- nacle Township, Burlington County, No- vember 4, 1915. (Courtesy of Herbert N. Morse.) Shear, C. L.: Cranberry Diseases. Bulletin 110, U. S. Bureau of Plant Industry, 1907. VEGETATION OF THE NEW JERSEY PINE-BARRENS 27 and by slow physic and chemic changes.* Such peat consists of the remains of bog mosses, as well as the leaves and stems of other bo^- inhabiting plants. Buried in this peat, as previously noted, are per- fectly preserved logs of white-cedar trees, which have been removed for their wood after the lapse of centuries. Huckleberry Picking. — Beneath the pine trees and in the swamps of New Jersey grow a number of species of huckleberries (Gaylussacia) and blueberries (Vaccinium). When the fruit of these bushes is ripe, the crop of berries is gathered, boxed, and crated for shipment to our large cities. This gives employment to many men, boys, women, and girls, now largely recruited from the foreigners who have immigrated to America. The blue huckleberry, Gaylussacia frondosa, is frequent in dry areas of the pine-barrens. It and the black huckleberry, Gaylussacia baccata, form the bulk of the huckleberry crop. The dwarf blueberries, Vaccinium pennsylvanicum, V. vacillans, yield fruit of great value when ripe in late June and early July. The swamp blueberry, Vaccinium corymbosum, frequent as a tall bush in swampy thickets in the pine- barrens, yields the largest and best-flavored berry, while the other tall blueberry, Vaccinium atrococcum, has a black fruit almost equally desirable. Experiments conducted by Coville indicate the practicability of cultivating the tall swamp blueberry by choosing soils with an acid reaction, so that in a few years the cultivation of this plant will be put on a commercial basis. t Drug Plants. — A list of the chief drug plants of the pine-barrens is given below, arranged according to the organs, or parts, of the plant used and the medicinal properties of each. The large drug houses still depend upon wild plants for part of their raw material, and the inhabi- tants of the districts in which these crude drug products are found make a livelihood collecting and curing the native plants for the large drug manufacturers. Roots Spurge (Euphorbia ipecacuanhae), diaphoretic, cathartic, emetic. Wild indigo (Baptisia tinctoria), stimulant, emetic, cathartic. Pitcher-plant (Sarracenia purpurea), unofficial. Goat's rue (Cracca (Tephrosia) virginiana), unofficial. Butterfly-weed (Asclepias tuberosa), unofficial. * Harshberger, J. W.: Peat Bogs and Peat. Old Penn Weekly, April 10, 1909, page 8; Bogs, their Nature and Origin. The Plant World, 12: 34, February, 1909; Parmelee, C. W., and McCourt, W. E.; A Report on the Peat Deposits of Northern New Jersey, Report N. J. State Geologist, 1905: 223-313. t Coville, Frederick v.: Experiments in Blueberry Culture. U. S. Bureau of Plant Industry, Bulletin 193: i-ioo. Plates 18, figures 31, 191 1. Also Nat'l Geogr. Mag., June, 1916. 28 vegetation of the new jersey pine-barrens Rhizomes Sweet-flag (Acorus calamus). Colic root (Aletris farinosa), tonic, emetic, purgative. Water-lily (Castalia (Nymphaea) odorata), demulcent, astringent. False sarsaparilla (Aralia nudicaulis), stimulant, diaphoretic, al- terative. Regal fern (Osmunda regalis), unofficial. Bark Swamp magnolia (Magnolia virginiana), diaphoretic, tonic, febri- fuge. White oak (Quercus alba), astringent. Black oak (Quercus velutina), astringent. Bayberry (Myrica carolinensis), acrid stimulant, sialagogue, errhine. Sassafras (Sassafras variifolium), stimulant, diaphoretic, alterative. Red maple (Acer rubrum), unofficial. Holly (Ilex opaca), unofficial. Alder (Alnus rugosa), unofficial. Wild cherry (Prunus serotina), stomachic and bitter tonic. Leaves Rattlesnake-plantain (Peramium pubescens), antivenine.* Bearberry (Arctostaphylos uva-ursi), astringent, tonic, diuretic, nephritic. Trailing arbutus (Epigaea repens), astringent, tonic. Laurel (Kalmia latifolia), astringent and poisonous. Holly (Ilex opaca), demulcent, tonic, emetic. Pipsissewa (Chimaphila umbellata), astringent, tonic, diuretic, nephritic. Wintergreen (Gaultheria procumbens), stimulant, astringent, diu- retic, emmenagogue. Sweet-fern (Comptonia asplenifolia), stimulant and astringent. Poison-ivy (Rhus radicans), irritant, rubefacient. Herbs Sundew (Drosera rotundifolia), pectoral, rubefacient. Golden-rod (Solidago odora), stimulant, carminative, diaphoretic. Horse-mint (Monarda punctata), carminative, stimulant, nervine, emmenagogue. Balsams and Oleoresins Pitch-pine (Pinus rigida) (turpentine), stimulant, diuretic, diapho- retic, astringent; (tar) stimulant, irritant, insecticide. * According to Captain John Carver, the Indians considered the juice of the leaves of the rattlesnake-plantain an absolute cure for rattlesnake poison. Applied externally and also swallowed, he says they were so sure of this fact that for a shilling they would let a rattlesnake bite them. This fact was contributed by Mr. J. L. Pennypacker, of Haddon- field. VEGETATION OF THE NEW JERSEY PINE-BARRENS 29 During the two hundred and fifty years in which the pine-barren region of New Jersey has been known to white men, a considerable num- ber of these drug plants have been gathered and utilized either as home remedies or else for shipment to Europe and to the larger American cities (notably Philadelphia), where they have been manufactured into useful drugs.* The revenue obtained by their sale has been a considerable asset to the herb gatherers of the New Jersey pine-barrens. Gathering of Greens and Flowers. — The income derived from the sale of greens for decoration, especially at Christmas time, and of wild flowers for bouquets, has been considerable to the people of the New Jersey pine forest. The collection of these materials has been so care- less and wasteful in past years that the supply of some of them is ex- hausted almost and all persons interested in the conservation of our vegetation and our natural resources should cry a halt. The following plants are those which have been gathered, according to the knowledge of the writerif Greens Ground-pine (Lycopodium obscurum). Pitch-pine (Pinus rigida). Greenbriar (Smilax laurifolia). Laurel (Kalmia latifolia). Mistletoe (Phoradendron flavescens). Holly (Ilex opaca). Inkberry (Ilex glabra). Cat-tail (Typha latifolia). Plume-grass (Erianthus saccharoides). Flowers Golden-club (Orontium aquaticum). Turkey-beard (Xerophyllum asphodeloides). Swamp-pink (Helonias buUata). Turk's-cap-lily (Lilium superbum). Moccasin-flower (Cypripedium acaule). Water-lily (Castalia (Nymphaea) odorata). Sweet-bay (Magnolia virginiana). Wild lupine (Lupinus perennis). Sweet pepperbush (Clethra alnifolia). Pink azalea (Azalea nudiflora). White azalea (Azalea viscosa). Sand-myrtle (Dendrium buxifolium). * Consult Henkel, Alice: Wild Medicinal Plants of the United States. Bureau of Plant Industry, Bulletin 89, Washington, 1906. t See Treat, Mary: Native Plants for Winter Decoration. Garden and Forest, vi: 141, March 29, 1893. 30 VEGETATION OF THE NEW JERSEY PINE-BARRENS Laurel (Kalmia latifolia). Arbutus (Epigaea repens). Butterfly-weed (Asclepias tuberosa). Buttonbush (Cephalanthus occidentalis). White boneset (Eupatorium album). Rough boneset (Eupatorium verbenaefolium). Sweet goldenrod (Solidago odora). The plants in the above list that have been most in demand until some of them are threatened with extinction are mistletoe (Phoradendron flavescens), water-lily (Nymphaea odorata), sweet-bay (Magnolia vir- giniana), holly (Ilex opaca), pink azalea (Azalea nudiflora), laurel (Kal- mia latifolia), and arbutus (Epigaea repens). It has been no uncommon sight to see negro women on the streets near the large markets of Phila- delphia selling large bouquets of wild flowers gathered in the pine-barrens of New Jersey, notably water-lilies, sweet magnolias, pink azaleas, laurel and arbutus, while at Christmas time boxes and crates of mistletoe, holly and laurel have been exposed for sale. All true lovers of our native vegetation look for some proper control of the ruthless destruction of our most beautiful trees, shrubs, and flowers for purely commercial purposes. The time has come to call a halt in the destruction of such natural re- sources. CHAPTER IV RESEARCH INVESTIGATION OF PINE-BARREN SOILS Considerable attention has been given in recent years to an analysis of the soil as related to the growth of native vegetation. This study has developed several lines of investigation. The Bureau of Soils of the United States Department of Agriculture has emphasized the physic analysis of soils as of importance in determining the character of the soil, and it has emphasized also the importance of the toxic and useful organic constituents of the soil, as in part an explanation of soil sterility or of soil fertility. The second class of investigators, as Hilgard and Shantz, have considered natural vegetation to be a valuable indicator of the capabilities of the land for crop production, while the third group of students have laid stress upon the importance of all the factors which influence the character of the soil, whether they be chemic, physic, toxic, or biologic. The writer has expressed his views* on this important matter as follows: " For many years the fertility of the soil was sought in the chemic substances which analysis proved to be essential to plants and which could be exhausted from the soil by the continual growth of a single crop upon it. To restore the fertility of the soil, it was necessary only to restore the ingredients necessary to keep a plant in a productive con- dition. Fertilizers were applied which were known to contain the most important materials of plant food and in an available form. Even to- day there are opposing camps of plant physiologists. One set holds to the principles, first clearly enunciated by Liebig, that the chemic con- dition of the soil is the most influential factor in the productivity of the garden or farm. The other group consider that the physic condition of the soil influences the tilth. This school teaches that all agricultural soils contain sufficient quantities of the essential mineral plant foods for many years to come. Recently a more advanced position has been taken by some students of the soil when they claim that the loss of fertility of many long cropped soils is due to the accumulation of toxic bodies, the accumulated excreta of plants that may have been grown without proper rotation. The true theory of soil fertility will probably be found to be * Harshberger, J. \V.: The Soil, a living Thing. Science, N. S. xxxiii: 741, May 12, 191 1. 3' 32 VEGETATION OF THE NEW JERSEY PINE-BARRENS one which will combine all of these theories with another one, which 1 believe must also be considered in reaching a satisfactory conclusion as to the relation existing between crops and the soil in which they grow. "The theory is one which considers that the soil is a living thing apart from its chemic or physic structure, that in the reaction between the living soil and the growing plant is the true explanation of soil fertility. A fertile soil is a live one. An infertile soil is a dead one. Contrast the soil which is filled with organic matter (humus) and in which numberless fungous, bacterial and protozoan organisms are at work, with a mass of clay or sand without such organic material and associated living organ- isms. The one soil is fertile, because the organisms in the soil react favorably upon each other; the other soil is infertile, because the organ- isms present in this soil are antagonistic. Recent investigation has pointed the way along which future research on soils must proceed. That the soil is the seat of activities of much importance to growing plants is proved by the presence of the nitrifying and denitrifying bac- teria, of the bacteria that produce the root nodules of the Leguminosae, of such organisms as Clostridium pastorianum, Bacillus mycoides, B. ellenhachensis, A^otobacier chroococcum, A. Vinelandii and the hyphae of numerous saprophytic fungi, various putrefactive bacteria, which perform their role in making the soil the fit habitation of the higher flowering plants, producing the tilth or 'Bodengare' of the Germans. So too earthworms, insect larvae, ants and burrowing animals assist in the task of aerating and mixing the surface layers of the soil. It is also evident that the production of toxic excretions by the roots of plants is undoubtedly a factor of importance in soil fertility. Following out a clue which the partial sterilization of the soil by chemicals or by steam gave, it was discovered that the bacteria which are useful in ammonia- making increased four-fold after such treatment, suggesting the presence in the soil of some agent which held them in check. After much pains- taking study it was discovered* that the soil contained a living proto- zoon (Pleurotricha), which preyed upon the useful organisms, and that the heat and chemicals either destroyed these larger unicellular animals, or inhibited their activity. It can be said, therefore, that the problem of fertility of the soil is largely a biologic one, as well as dependent upon the physic, chemic and toxic condition of the surface layers." The attempt has been made in the investigation of the pine-barren soils to make the study from an ecologic point of view. Nine stations were chosen in different parts of the region in order that the study of the pine-barren soils might be made as comprehensive as possible. As the * Hall, A. D.: The Soil as a Battleground, Harper's Magazine, October, 1910, pp. 680-687. VEGETATION OF THE NEW JERSEY PINE-BARRENS 33 sand of central New Jersey was derived originally from oceanic supplies, the first station (I) where soil samples were taken was the beach and the dunes at Belmar. The second station (11) was chosen at Como, where the pine-barren forest approaches the sea. This forest may be termed the coastal pine forest. Station 111 was located in the pine forest about 1.6 kilometers (i mile) east of Lakehurst, and the other stations were located at Warren Grove (IV), at Shamong (Chatsworth) (V), at Atco (VI), at Sumner (VI I), at Clementon (VI 1 1), and in the de- ciduous forest at Clementon (IX). Some 28 samples were taken from the 9 different stations and some of the generalizations reached in this study are based on a study of these 28 samples representing a wide diversity of conditions of pine-barren soils. Mechanic Analysis of Soils Nineteen samples of soil were sent to the Bureau of Soils in Washington for mechanic analysis. The bureau very considerately undertook the task of analyzing these samples, and after a lapse of considerable time reported, under date of January 10, 191 1. In the following table the analysis of 16 samples is given: Med- Very Serial Fine Coarse Sand ium Fine Sand Fine Silt Clay Num- Note GiiAVEL Sand Sand 0.05 to 0.005 ber, Book 2 TO I I TO 0.5 TO 0.25 O.I TO 0.005 TO 0 U. S. Num- Locality Depth MM. 0-5 M.M. 0.25 TO o.i 0.05 MM. .MM. Bureau ber OF mm. MM. MM. OF Sample Soils P. ct. P. ct. P. ct. P. ct. P. Ct. P. ct. P. ct. 23821 I Belmar Beach Sand 8.5 59-2 28.0 4-1 O.l o.i 0.0 23822 2 Surface Dune 1.2 47-4 39-3 9-7 0.7 I.O 0.4 23823 3 Dune 30 cm. 2.1 59-7 29.1 6.3 1-5 0.6 0.4 23824 4 Como Surface Soil 3-1 20.5 26.S 34-9 8.0 S-i 1-3 2382s 5 10 cm. 3-2 23-4 27.3 29.4 9.9 5-3 1-5 23826 6 24 cm. 9.8 28.2 27.2 21.6 7-7 4-3 l.O 23827 7 55 cm. 3-2 21. 1 30.4 27.6 5-8 6.1 S-9 23828 8 Lake- hurst Surface Soil 5-3 22.3 18.6 43-1 4.1 4.1 1.6 23829 9 Lake- hurst 15 cm. 4-9 20.0 21. 1 42.6 7.4 2.5 1.2 23830 10 Lake- hurst 50 cm. II. 8 18.6 19.4 35-4 5.6 4.4 4-5 23831 1 1 Upper Plains Surface Soil 1.2 13.8 28.8 41-3 9-5 i-3 2.0 23832 12 Upper Plains 15 cm. 2.5 15-4 25.8 39-4 9.1 4.8 2.8 23833 13 L'pper Plains 40 cm. 1.2 II. 2 25.4 37-8 10.6 10. 0 3.8 23837 17 Shamong Surface Soil 25 cm. 2.9 28.3 23.4 35-3 6.5 1.8 i.S 23838 18 Shamong 3-2 18.9 23.0 43.0 9.1 1.4 1-3 23839 19 Shamong 50 cm. 3-0 19.4 22.7 40.2 8.0 2.6 3.6 34 VEGETATION OF THE NEW JERSEY PINE-BARRENS It will be noted that fine gravel, where the particles range from 2 to i millimeters, is found in the coarse beach sand, the middle layer (24 centi- meters deep) of the soil at Como (9.8 per cent.), at 50 centimeters deep in the soil at Lakehurst (i 1.8 per cent.), 15 centimeters deep in the soil from the Upper Plains (2.5 per cent.), and at 25 centimeters deep in the soil from the pine woods at Shamong (3.2 per cent.). The soils .in gen- eral from the five stations given above range from coarse to fine sand (i mm. to o.i mm.). Small percentages of silt not exceeding 6.1 per cent., with one exception, 10 per cent., in the soil 40 centimeters below the surface in the Upper Plains, are noted. The percentage of clay in- creases in the soils from the inland stations, being uniformly larger in the three samples from the Upper Plains. In general, therefore, it may be stated that the soils of New Jersey pine-barrens, from the surface to a depth of 55 centimeters, are sandy with slight percentages of silt and clay. MiNERALOGIC ANALYSIS OF SoiLS A mineralogic examination of 19 samples of soils furnished by me to the U. S. Bureau of Soils is as follows: The sandy soils show approximately the same mineral composition, varying somewhat in the percentage of the same. They are generally poor in minerals other than quartz, i. e., 3 to 5 per cent, and less. The feldspar content is very low and generally highly altered. The minerals commonly found, in the order of their occurrence, are: zircon, rutile, sillimanite, tourmaline, feldspar, and hornblende. Magnetite and ilmenite are present in some quantity. Augite, epidote, titanite, and chromite are present occasionally. The quartz grains, zircon and rutile crystals, indicate by their rounded ap- pearance a great amount of attrition. There are no distinctive differ- ences in the samples. Chemic Analysis of Soils Through the kindness of Dr. Daniel L. Wallace, of the Department of Chemistry of the University of Pennsylvania, the following partial anal- yses of surface, middle and subsoil layers are presented. The surface, middle and subsoil materials submitted to analysis were obtained by mixing soils from the different localities mentioned above and from all three levels: Nitrogen Potash* (KjO) Phosphorus (P2O5) Surface layers . Middle layers. Subsoil layers. 0.24 p. c. 0.05 p. c. 0.06 p. c. 0.53 p. c. 0.24 p. c. 0.35 p. c. 0.07 p. c. o.io p. c. 0.06 p. c. * The original analysis for potash gave the percentages in terms of K2CO3, but to har- monize them with other analytic data they were converted into terms of K2O. In terms of K2CO3 the percentages were 0.78, 0.35, 0.52. VEGETATION OF THE NEW JERSEY PINE-BARRENS 35 It will be noted that the surface layers are richer in nitrogen. As to potash, the surface soils are richest, the middle layers less so, and the subsoils have a higher percentage than the middle layers. The middle layers are richer in phosphorus than either the surface or subsoil layer. Contrasting this with a high class soil such as Hilgard (Soils, 331) gives from Janesville, Wisconsin, with potash 0.59 per cent., and phosphoric acid (P205),o.o6, where the phosphoric acid is only just above the lower limit of sufficiency, we discover that the pine-barren soils of New Jer- sey, as far as phosphorus is concerned, are richer in this ingredient than the rich, friable loam of Wisconsin. The percentage of potash in the New Jersey soils is less than the percentage of that element in the Wisconsin soil. Hilgard, on page 352 of his book on Soils, gives for the pinewoodsof Mississippi potash (K2O), 0.26 per cent., and for the pine flats, 0.06 per cent., and phosphoric acid (P2O5), 0.03 per cent. For high-class pinelands of Florida he gives potash, o. 19 per cent., and phosphoric acid, 0.1 1 per cent. In comparison with these Florida soils the potash con- tent of the New Jersey is higher, but as regards phosphorus, there is a slight difference in favor of the Florida soils. The nitrogen content is, however, low as compared with the limits set by Hilgard (358) of 5 per cent, for humid regions and i per cent, for arid regions. Hence the New Jersey soils ought to respond to the use of nitrogen. Experiments with Pine-Barren Soils The results of a series of experiments upon pine-barren soils are given in the following table for 28 samples from 9 diverse localities in the region. In order to determine the water content of each sample, 500 c.c. of each kind of soil was taken and carefully weighed in the fresh condition. In each case the measure was filled heaping full, and the soil was shaken in order to compact it before the sample was emptied on to a piece of paper on the pan of the balance. After weighing the samples were spread upon newspapers and thoroughly air-dried before being carefully weighed again. The difference between the first weight and the second weight gave the weight of the water lost during the process of air-drying. The rate of percolation was estimated by taking a wide glass tube, 4.2 cm. wide and 49.8 cm. long, and marking upon it circular lines 5 cm. apart up to 30 cm. (Fig. 11). The lines were marked 0,5 cm., 10 cm., 15 cm., 20 cm., 25 cm., 30 cm. A piece of wet cheese-cloth was tied to the bot- tom of this glass cyHnder, 2.3 cm. below the 30 cm. mark on the tube, which was filled with soil some distance above the zero (o) mark, 17 cm. below the top rim of the glass cylinder. The soil was shaken into place. 36 VEGETATION OF THE NEW JERSEY PINE-BARRENS but not tamped. The location of the zero point below the top of the tube was purposely made, so that the sand above it would equalize the flow of water before it reached the zero line of the tube. The rate of flow from the zero line to the line marked 30 cm. was noted by means of a stop-watch reading to seconds (Fig. 11). The time that it took for the different columns of water to reach the different 5 cm. levels were noted by the stop-watch. As the 30 cm. mark was 2.3 cm. above the cheese-cloth, the total time is not a sum of all the time intervals, but is the time that it took for the water to pass from 30 cm. mark to the bot- tom of the tube, as indicated by the first drop falling to the catch-vessel Figure 1 1 Apparatus used in conducting experiments with pine-barren soils. Three glass cylinders are shown with tripod supports, cheese-cloth to hold soil in place, and receptacles to receive drainage waters, December 7, 19 10. The stop-watch is held in the left hand of the author. below, plus the time that it took to pass through the 30 cm. column of soil. As 500 c.c. of water were used for each percolation experiment, the amount of percolation water was determined by measuring the amount of water which passed through the column of sand into the receptacle beneath after the flow of the water had ceased. Deducting the water of percolation from the original 500 c.c. used, the difference indicates the amount of water in volume actually retained by the soil. These figures are given also in the table. As these samples were taken at different seasons of the year, the water content of the different soils, as indicated by the loss of weight of the air- dried sample of soil, is of course different. For example, it was 124 VEGETATION OF THE NEW JERSEY PINE-BARRENS 37 grams in the surface soil from the pine woods one mile east of Lakehurst, 128 grams from the sample taken 40 cm. below the surface in the de- ciduous woods one-quarter mile west of Clementon, and 209 grams in the sample taken 61 cm. below the surface in the same locality. The lowest was 5 grams from the beach sand gathered at Belmar. The two extremes, 5 grams for the beach sand and 128 grams and 209 grams for the subsoil upon which grew the deciduous forest, is of interest. For the pine woods, taking the mean for all of the samples of the surface layer, it stood 73 grams; for the root-containing layer it was 35 grams; for the deeper samples in the rootless subsoil it was 33.4 grams. It is important to give a detailed description of the stations where the 28 soil samples were taken before considering the table of percolation experiments. Station I. — Belmar. Sample i. — Upper beach sand (surface). Sample 2. — Dune sand (surface) between Ammophila arenaria and Solidago sempervirens. Sample 3. — Dune sand, 30 cm. below the surface, with growth of Ammophila arenaria, Cassia chamaecrista, Solidago sempervirens, and Strophostyles helvola. Station II. — Pine woods along north shore of Como Lake. Sample 4. — Surface, or humous, layer immediately beneath the cover- ing of pine needles, which were removed. Sample 5. — Sandy soil, 10 cm. below the surface. Sample 6. — Sandy soil, 24 cm. below the surface and directly under- neath the horizontal pine roots. Sample 7. — Fine red-gravel layer, 55 cm. below the surface. Station 111. — Pine woods, 1.6 kilometers (1 mile) east of Lakehurst. Position of Quadrat II. Sample 8. — Center of Quadrat II. Surface soil. Sample 9. — Soil 15 cm. below the surface. Sample 10. — Soil 50 cm. below the surface. Station IV. — South edge of Upper Plains at Warren Grove, near George Cranmer's farm. Sample 1 1 . — Surface soil. Here grew Pinus rigida, Dendrium buxi- folium, Gaylussacia resinosa, Ilex glabra, Pyxidanthera barbulata, Quer- cus ilicifolia, Tephrosia (Cracca) virginiana. Sample 12. — Soil 15 cm. below the surface. Sample 13. — Soil 40 cm. below the surface. Station V. — Pine-barrens at Shamong (Chatsworth). Sample 17. — Surface, or humous, layer beneath the tall pine trees. Sample 18. — Soil 25 cm. below the surface. Sample 19. — Soil taken 50 cm. below the surface. 38 VEGETATION OF THE NEW JERSEY PINE-BARRENS Station VI. — Pine woods at Atco, November 8, 1910. Sample 20. — Surface, or humous, soil. Sample 21. — Soil 10 to 15 cm. below the surface in the root layer. Sample 22. — Soil 50 to 60 cm., below the surface. This soil was wet and cold, and was taken probably near the edge of a deciduous swamp. Station VII. — Pine forest at Sumner. The humous layer was 20 cm. deep, of dark-grayish sand. Next came a white, sandy layer, 40 cm. thick, with reddish-yellow sand beneath. The component vegetation here consisted of Pinus rigida, Gaylussacia resinosa, Gaultheria procumbens, Kalmia angustifolia, K. latifolia, Pterid- ium aquilinum, Quercus ilicifolia, Q. prinus. Sample 23. — Humous layer down to 15 cm. Sample 24. — Sandy soil, gray in color, 45 cm. below the surface. Sample 25. — Subsoil reddish-yellow sand, 61 cm. below the surface. Station VIII. — A deciduous forest, 0.4 kilometer (one-quarter mile) west of Clementon, with Liquidambar styraciflua, Liriodendron tulipifera, Quercus alba, Q. velutina, Euonymus americanus, and Viburnum dentatum. Thetopsoil was full of humus, about 30 cm. thick, followed by a thin layer 25 cm. thick, succeeded by a coarser sand saturated with ground-water. Sample 26. — Surface soil below leaf-covering down to 10 cm. Sample 27. — Soil 40 cm. below the surface. Sample 28. — Soil 61 cm. below the surface, with water running into hole as fast as the sand was dug out. Station IX. — Deciduous woods on a sloping hillside at Clementon. The sandy ridge or hill with sand at least 6.4 meters (21 feet) thick was covered with such trees as: Acer rubrum, Cornus florida, Dios- pyros virginiana, Fagus grandifolia. Ilex opaca, Liquidambar styra- ciflua, Pinus rigida, Quercus alba, Q. velutina. Sassafras varii- folium, such shrubs as Kalmia latifolia, Smilax rotundifolia. Vi- burnum dentatum, and the herbs Epigaea repens, Maianthemum bifolia and Mitchella repens. Sample 29. — Top soil below forest litter down to 12 cm. June 9, 191 1 . Sample 30. — Soil 23 cm. below the surface. Sample 31. — Soil 38 cm. below the surface. The rate of percolation, as shown by the table and by the graphic curve, varied greatly in the different soil samples. The curves for the percolation through the surface soil samples show what diverse condi- tions exist in the humous layer in the pine-barren region. The curves I and 2 for beach and dune sands show that the passage of water through them was extremely rapid. The curves of samples 4 and 8 conform nearly with those of the beach sand and show that the perco- lation of water through the surface soil in the pine woods at Como and Lakehurst is rapid. The curves for samples 17 and 18, although steeper and more abrupt, indicate that the rate of percolation is slower, but still VEGETATION OF THE NEW JERSEY PINE-BARRENS 39 rapid enough, so that the surface soils at Shamong and at Atco can be called pervious. Not so with the steep and abrupt curves of samples 23, 26, and 29 (Fig. 12), for the pine woods at Sumner, at the western edge of the pine-barrens, at Clementon. Here the cates a slow rate of per- curves which represent through the root-holding we discover that with the of sample 3, that of dune curves 6, 9, 12, 18, 21, 30 conditions existing in the uniform throughout the of the surface soil, as might abrupt and lengthy curves the soil at Sumner, 55 cm. deciduous forest soil 40 and for the deciduous woods steepness of the curves indi- colation. If we examine the the rate of passage of water layers of the pine-barren soils, exception of the very rapid rate sand 30 cm. below the surface, approximate, showing that the root-holding layer are more region than are the conditions be expected. The extremely 24 and 27 respectively for below the surface, and for the cm. below the surface, 0.4 kilo- meter (one-quarter mile) west of Clementon, indicate that these soils are extremely im- pervious (Fig. 12). The series Figure 12 Curves showing rate of percolation of water through various soils. A, Surface soil; B, root-holding layer; C, rootless subsoil. of curves for the rootless subsoil, with the exception of the extremely pervious subsoil 55 cm. below the surface in the pine woods at Como, numbered 7, are also fairly uniform. Curves 19, 22, 28 form one group, 40 VEGETATION OF THE NEW JERSEY PINE-BARRENS and curves lo, 13, 25, 31 form another group, the subsoil of the second group being less open and with a slower rate of percolation than the first group. Now as the soil samples from the different stations were taken from the same hole, but at different depths, we must connect the different curves in a series by way of determining the rate of percolation under natural conditions of soil environment where the humous layer is super- imposed on the root-holding layer, and it in turn on the rootless subsoil. For Station II, curves 4, 6, 7 form a series; for Station III, curves 8, 9. 10 are a series; for Station IV, curves 11, 12, 13 form a series; for Station V, curves 17, 18, 19 should be grouped; for Station VI, that at Atco, curves 20, 21,22 arrange themselves in a series; for Station VII, curves 23, 24, 25 can be grouped; for Station VIII group the curves 26, 27, 28, and for Station IX, curves 29, 30, 31 form a series (Fig. 12). To sum up as to the rate of percolation of water through the soils at these nine stations: Station II.- — Percolation rapid through the surface layer, slower through the root-holding layer, rapid through the subsoil. Station III. — Percolation rapid through the surface soil, not so rapid through the second soil layer, very slow through the subsoil. Station IV. — Percolation was slow through the surface soil, more rapid through the root-holding layer, and again slower through the sub- soil. Station V.— Percolation through the three layers of soil from the pine woods at Shamong was fairly uniform, being slightly retarded in the middle layer. Station VI. — The soil at Atco was relatively pervious, especially the surface layer. Station VI 1. — The soil of the pine woods at Sumner was relatively impervious, especially the middle layer. Station VIII. — The soil of the deciduous forest, 0.4 kilometer (one- quarter mile) west of Clementon, was as indicated by the curves 26, 27, very impervious, but the subsoil, indicated by curve 28, was more open and porous than the upper layers. Station IX. — The rate of percolation through the soil layers of the hillside deciduous forest was much slower than through the pine-barren soils, as indicated by the steepness of curves 29, 30, 31. If we may be permitted to generalize on this series of experiments, it seems obvious that we have the dune sands at one extreme and the soils of the deciduous forest at the other extreme. The dune sands are open and pervious to Vv^ater. The soils of the deciduous forest allow the water Deciduous Forest (Hillside) at Clementon IX 3.1EJ -jng Axopq -UID 8? l!OS ■0 W M 0 li CO ri 0 0 -t 1- -. 0 10 10 10 0 0 0 1 t^ " OC 10 10 0 0 aauj -jng A\opq -uia fz pog 0 n oroir«oinioo 0 0 aaj5n jKaq .wopq pog aaEjang Ov r- t^oioinoooio \n in Deciduous Forest 0.4 Kilometer West of Clementon VIII aa-dj -jng A\opq 'luo 19 pog CO ^ -rov>n>ooir.oo 0 0 0 Oi 0 M too N 00 " Ov >- 00 ON MWPir^N ajBj -jng ,wopq -uia ot pog r^ r^ C>00 OIOIOOOO -t 0 01 0 (N 0 10 TfsO Ol r^ \ri Tt ^anrj j-eaq A\opq pog aaEjang 0 CO -ftminoOOO -0 't Pine Forest AT Sumner VII ajBj -jng Mopq -uia 19 pog lo 10 "Tj-r~OONOO 0 0 « t^ t M in 0 vO 10 ro t~ ~> aoEj -jng A\oiaq -ujd SS pog •* 0 ^oinomoini/i 0 0 vo 0 «tiot^a M n ja.\Eq snouinj-i JO ^OBjjng ~5 00 OCOI^IOIDI/^INO N CO - l/:0 OloOmNfOtN ro \0 Pine Forest AT Atco VI aaBj -jng A\opq -uia o£ pog s r~ 0000000100 0 -3- t^ 0 0 ►"• 'i"0 c^ ^ t^ 0 0 00 M h-i (N M aoBj -jng A\opq -lua £1 pog ?; 0^ 0 O>oo irj ro 10 w m 0 0 ro 00 l/^ 0) l/)00 M -0 t^ ^ IT) Tf *-i ^ M N jBABq snouinH jo aaejjng 0 " 1-100 1/110 10 0 0 0 M rOOifOlONlOOlO 10 lO •rf •-' 04 M f^ t^ 0 1000 0 '-0 0 0 M W KH M M aaBj -jng A\opq -010 £z pog 00 r^ Oi-iOiONOioio 0 0 ■t «fv:Nioo>'tO"t "i" 0 0 0 M w n (N CO^IOOIOOOO 10 10 10 rOcjNO0rO^'-'N aanj -jng A\opq -ma £1 pog 0 CO ooOootl^OioO 10 10 0 00 01 w TtO 0 ro r^ ro O 0 10 H, « « n 01 isajoj auij UI pog aaejjng 00 00 ^Tl-tt^vOlO .0 10 10 0 0001 MOO.tlOTt 0 t-H N <-< Pine Forest, North Shore of CoMO Lake II aaBjjng .wopq ■uia ££ 'pog pABjf) pa^ t^ t~ ro^iOvOOrOOO lo lo hH t^ .,:}• M ro rtsO r^ ro sO sO 10 ~5 " aaEjjng .'k\opq ■uia tz 'pog ApUBg OiorOOOOlOO 0 0 t^ 01 T)-M Tfa lo n 0 t^ ro 0 0 w 01 ro 01 01 aa-Ejjng A\opq •ma 01 'pog .^puBg . 0 Ot^OOOOOio 0 0 r^ roro ojTj-r^win 00 01 saipaax au'd JO JaABq qqsauaq jaXBq snomnn "* -t r^i^.sOocO'-'io 0 0 0 " t . -H ro 100 ro 1^ ■* t ro " Beach Sand, Belmar I aaBjjng A\opq •UJD of 'puBg auna '^ r^ 0000001010100 0 0 0 fOoi MMfO-^io fo r^ jaABq aaejjng 'pueg aunQ . r~ 0 t^ . r-o 0 ro (^ lo 10 10 10 . f-tro-^io-^oo 00 rO M ja.iEq aaBjjng 'qaBag jaddQ - 0 -tiO<*"00". 10 10 00 00 "0)rO-!f. tt 00 ~; -H a a to a. a 0 W The lines 0 to 30 cm. refer to the intervals of the columns of soils, and the figures in the corresponding columns are sec- onds of time. 500 C.c. Fresh Soil. Weight in grams . 500 C.C. Air-Dried Soil. Weight in J3 'v si l-J c E 1 E c 1 c 5 1- E c 1 E c 1 50 25-30 cm Amount in c.c. of Percolated Water Amount in c.c. of Retained Water . 4" 42 VEGETATION OF THE NEW JERSEY PINE-BARRENS to pass less freely, in other words, they have a greater capacity for holding water. The pine-barren soils are sandy and pervious and may be said to occupy an intermediate position between the two extremes mentioned above. In all probability the deciduous forests exist on the less pervious soils, because they can exist solely in a soil which, by its water-holding capacity, can supply the water given off in transpiration from the enor- mous surface of the broad leaves. The pine, as a true xerophyte, occupies those soils which are more pervious and which show a more rapid rate of percolation, while the dune sands, being entirely without silt and clay, are covered by an arenicolous vegetation distinct from that of the pine forest. Geographically, the pine-barrens occupy an inter- mediate position between the strand vegetation on the east coast of New Jersey and the deciduous forests in the west. Experiments with Capillary Rise of Water A few experiments were conducted on the capillary rise of water in air- dried soil, but on account of the length of time that it took to complete the tests, they were discontinued. The experiments consisted in im- mersing the glass tube containing the soil in a bottle of water and noting the rate of capillary rise. The following figures are given, and the sample numbers are the same as given in the preceding pages. The numbers on the glass tubes are reversed, reading from bottom to top. The time is given in seconds for uniformity. The dry soil was shaken together into the tubes. Sample 2 : Sample 5: 30-25 cm.= II seconds 30-25 cm.= 25 25-20 " = 85 25-20 " = 62 20-15 " = ''637 20-15 " = 270 Sample 3 : 15-10 " = 1,050 30-25 cm.= 18 seconds Sample 6: 25-20 " = 135 30-25 cm.= 105 20-15 " = 1,920 25-20 " = 265 15-10 " =23,880 20-15 " = 600 Sample 4: 15-10 " = 1,703 30-25 cm.= 60 seconds Sample 7: 25-20 " = 210 30-25 cm. = missed 20-15 " = 706 25-20 " = 30 15-10 " = 3,840 20-15 " = 240 15-10 " = 1,620 seconds seconds start seconds Large stones depress the rise of the water on the side of the glass tube where they occur. As the data for the capillary rise of the water in the column of sand are not complete, no graphic curves have been drawn. VEGETATION OF THE NEW JERSEY PINE-BARRENS 43 However, as the time intervals for the rise of water between the suc- cessive 5 cm. are large, the curves would be sharp and steep for each of the soil samples. The rise of water from a saturated subsoil through surface layers of dry sand during periods of drought must be extremely slow, and that the lack of this water is felt by plants having superficial root systems is reflected in the xerophytic structure of these plants. The deep-rooted oaks and pines are more independent of the surface supplies of water, as their root systems usually penetrate into the wetter subsoil, while the superficially rooted plants, cut off from the deeper supplies of water, are dependent on the occasional light showers and are able to survive because of the structures which enable them to reduce their water loss. Retention of Water by Pine-Barren Soils As the water which drains through a soil cannot be of any permanent value to the vegetation found on that soil, it is important to ascertain the capacity of a soil to retain the water which falls as rain. The re- tentive capacity of the different soil samples is given on the last hori- zontal line of the foregoing table. A graphic representation of that re- tentive capacity will enable us to interpret the phenomenon (Fig. 13). It will be noted from the accompanying graph that the dune sands (Sta- tion 1) are at the extreme of minimum retentivity (145, 185, 170 c.c. respectively, out of 500 c.c. of water used), and that the deciduous forest soils at Clementon (Station IX) are at the other extreme of maximum retentivity of water (325, 330, 300 c.c). The soils of the pine-barrens (Stations 11, 111, V, VI, VII) and Upper Plains (Station IV) stand mid- way (Fig. 13). With the exception of the coastal pine forest, the soil of which is more or less influenced by proximity to the dune sands, in none of the pine-barren soils does the amount of water retained by the soil out of 500 c.c. used fall below 200 c.c. The surface soil of the coastal pine-barrens retains only 170 c.c. of water, and the red gravel soil, 55 cm. below the surface, only 165 c.c. In many cases, chiefly for the sur- face soils of the pine-barrens, the number of cubic centimeters of water retained by the soil out of 500 c.c. used reads 245, 230, 250, 268, 268; for the middle soil layers the numbers are 230, 265, 235, 260, 230, 230; and for the subsoil they stand 200, 225, 235, 204, 230. The arithmetic mean of the figures expressing the amount of water retained by the dune sand is 166.66 c.c, or 33.33 per cent, of the 500 c.c. originally used in the experiments. The mean for the Upper Plains is 230, or 46 per cent.; the mean for the 16 pine-barren determinations is 229.37, almost that of the Upper Plains, or 45.87 per cent., while that of the deciduous forests 44 VEGETATION OF THE NEW JERSEY PINE-BARRENS is 280.83, or 56.16 per cent. It should be noted that the water-retaining capacity of the soils from the deciduous forest at Station VI 11 is not much different from that of the pine-barren soils. This may be ac- counted for by the close geographic position of the deciduous forest at Clementon near the western edge of the pine-barren region (Fig. 12). Figure 13 Graph representing the water-retentiveness of pine-barren soils. From this study we conclude that the dune soils have the fastest rate of percolation and the least retentive character. The soils of the decidu- ous forest have the slowest rate of percolation and the greatest reten- tiveness, while the pine-barren soils have an intermediate rate of per- colation and soils of fair retentiveness (Fig. 13). vegetation of the new jersey pine-barrens 45 Wilting Coefficient of Pine-Barren Soils The higher plants obtain from the soil by means of their roots the water upon which their life and growth depend. It is taken up in the main by the root-hair cells near the growing tips of the roots. For each type of soil there is a definite limit of moisture content, below which the water cannot be absorbed by plants in sufficient quantity to supply their needs. This limit varies with the moisture-holding capacity of the soil, being high in clay and low in sand. The plants wilt when this limit is reached, and remain wilted until the soils get additional moisture. Hence this limiting moisture content has been called the "wilting coefficient." It is believed commonly that some plants are better able than others to extract the water from relatively dry soil. It would seem almost self- evident that those plants which thrive in regions where drought occurs must be able to lower the water content of the soil to a point far below that at which the roots of ordinary plants cease to take up moisture. Nevertheless the experiments of Briggs and Shantz* have shown that there is little difference in this respect, wilting in all cases taking place at practically the same limit of moisture content in a different soil. The formula to determine the wilting coefficient of a soil when the water-holding capacity is known has been determined by Briggs and Shantz to be: „,., . ^^ . moisture-holding capacitv — 21 Wilting coefficient = — - — '—r — 2.90 (i ± 0.021). The moisture-holding capacity is the percentage of water a soil can retain in opposition to the force of gravity when free drainage is pro- vided. This percentage can be determined readily for the sample soils of the pine-barren region studied and described in the foregoing pages. In order to reduce the mathematic calculations to a minimum, the wilt- ing coefficient will be determined for dune sands, Upper Plains soils, pine-barren soils, and deciduous forest soils by striking a mean of all the observations made on these four types of soils. The moisture-holding capacities derived from the arithmetic means of all the determinations made for the four types of soils are as follows: Dune Sands 33-33 per cent. Pine-Barren Soils 45-87 " Upper Plains Soils 46.00 " " Deciduous Forest Soils 56.16 " * Briggs, L. J., and Shantz, H. L.: The Wilting Coefficient for Different Plants and its Indirect Determination, Bulletin 230, Bureau of Plant Industry, 1912. 46 VEGETATION OF THE NEW JERSEY PINE-BARRENS From these figures by substitution in the above formula we obtain the wilting coefficient of the four types of soils. Two values are obtained for each coefficient if we use the plus or the minus quantity in the last parenthesis of the formula. The wilting coefficients obtained in this way are: „ „ , / 4.1655 for + value. Dune Sands I 4.3569 " - " „. „ c- ■^ \ 8.0641 for + value. Pine-Barren Sous | g S\r << _ Upper Plains Soils 8.4635 for + value. 8.8339 " - " Deciduous Forest Soils | ^2.1628 for + value. I 12.3674 — According to the above computation, the pine-barren and the plains soils stand intermediate between the dune sands and the deciduous forest soils as to their wilting coefficients. In order to determine whether the above formula had been applied correctly, a letter was addressed to Dr. H. L. Shantz. His reply, under date of December 28, 191 2, makes clear some additional unpublished facts: "In applying the formula for the determination of the wilting coefficient from the moisture-holding capacity, or any of the formulae given in Bulletin 230, it is best for ordinary purposes to ignore entirely the quantity within the parentheses, since it is only an expression of the uncertainty of the correlation from our results. The amount of varia- tion which it will give in the values can be estimated as you have done, but this can be regarded by no means as the limits of error, when the wilting capacity is determined from the moisture-holding capacity. In handling these results 1 should simply deduct 21 from the moisture- holding capacity and divide the remainder by 2.9, and in case you wish to determine the amount of uncertainty with respect to a particular soil sample, the dividend would become in this case either 2.96 or 2.84, as the case might be. The most important thing in connection with the moisture-holding capacity is that the determinations be made under the conditions which were observed in our determinations, that is, in a column of soil one centimeter in height. In making this determination a great variation in values can be obtained, dependent upon the amount of shaking or jarring, and the determination is by no means a very accurate one." Adopting, as we have done previously, Shantz's second alternative, the vvilting coefficients of the four types of soils stand approximately as VEGETATION OF THE NEW JERSEY PINE-BARRENS 47 given above. If we use 2.9 as the divisor and carry our calculation to two decimal places, the numbers corresponding to the wilting coefficients stand: Dune Sands 4.25 Pine-Barren Soils 8.57 Upper Plains Soils 8.62 Deciduous Forest Soils 12.12 It will be noted that there is a discrepancy between the values of the •wilting coefficients derived by one or the other of these methods of de- termination, but the last figures may be taken as approximate expres- sions of the wilting coefficients of the different soils studied. Finally it ought to be stated that that portion of the soil moisture which is avail- able for plant growth is represented by the difference between the actual water content and the wilting coefficient. The latter determination is consequently essential in any critical study of plant growth to soil mois- ture. The Moisture Equivalent Composite samples of pine-barren soils were sent to Drs. Shantz and Briggs, who examined the soils and reported as follows: "The moisture equivalent of the composite sample of the surface, or humous layers of pine-barren soils numbered 4, 8, 17, 20 is 1 1 .3; for the middle soil layers numbered 5, 6, 9, 18, 21, it is 2.3, and for the subsoil layers numbered 7, 10, 19, 22, it is 4.4. The moisture equivalent of the soil is the per- centage of water which it can retain in opposition to a centrifugal force 1000 times that of gravity. In making the determinations which are expressed as percentages of the dry weight of the soil used the soils are placed in perforated cups and moistened with an amount of water in excess of the quantity they can hold in opposition to the centrifugal force. After standing twenty-four hours the cups are placed in a cen- trifugal machine, which is operated at a constant speed so chosen as to exert a force 1000 times that of gravity upon the soil moisture. This method provides a means of determining and comparing the retentive- ness of different soils for moisture when acted upon by a definite force, which is measured in absolute terms and is reproducible within narrow limits." CHAPTER V PHYTOGEOGRAPHIC FORMATIONS Considerable space has been given in the foregoing pages to a considera- tion of the geography, physiography, settlement, and industries of the New Jersey pine-barren region, with the object of assembhng the facts known about these subjects of general interest to those who have visited the region. The preceding account has dealt with those phases of the life of the people of south New Jersey which have been influenced by the vegetation, and thus it has been an ethnobotanic survey as far as that has been accomplished. The description of the pine-barren vegetation, as a whole, will follow, and the subdivisions of this vegetation will be dealt with under the head of the different and easily characterized plant formations. These formations have been distinguished in a general and unscientific way by the people who live in the pine-barrens with a keen appreciation of the fundamental differences in the native vegeta- tion, and this seems to be the case in other regions, as Graebner has emphasized in "Die Heide Norddeutschlands," and as I have detailed in my monograph on the Vegetation of South Florida.* It is left for the trained botanist to interpret these general impressions and to trans- late them into a scientific description based upon phytogeographic principles. When this is done, the phytogeographer recognizes nine natural plant formations and four that are due to fire, repeated cutting of the forest, or to the conversion of cedar swamps and bogs into areas suitable for cranberry culture. Classified in the order of their importance they are: Pine-Barren Formation. Cedar Swamp Formation. Deciduous Swamp Formation. Savanna Formation. Marsh Formation. Pond Formation. River Bank Formation. Bog Formation. Plains Formation. Cranberry Bog Formation. SuccEssiONAL j Scrub-Oak Formation. Plant Formations ] Oak Coppice Formation. Mixed Pine-Oak Formation. Natural Plant Formations * Graebner, Paul: Die Heide Norddeutschlands. Die Vegetation der Erde, v: 14; Leipzig, 1901; Harshberger, J. W.: The Vegetation of South Florida. Transactions Wagner Free Institute Science, vii, Pt. 3: 146, 1914. 48 VEGETATION OF THE NEW JERSEY PINE-BARRENS 49 The relation of these different formations to each other, as to their possible derivation in an ascending series toward a climax condition, may be represented in a graphic way (Fig. 14), where the comparative age of the formation is shown by its distance above the lowest line and the probable succession by means of dotted lines and arrows. As an illustration of the successions which the above diagram (Fig. 14) shows, the series beginning with the typic pine forest and ending with the mixed pine-oak formation may be chosen. When a pine forest is destroyed by fire, or the trees removed in lumbering operations, it may be succeeded by a scrub-oak thicket. This in turn is replaced by a coppice of taller Interrelation of the Pine-Barren Plant Formations. Pine Barrens. fjjygj. Bank. ^Deciduous Swamp. .^ ^Pond >Cedar Swamp. -^ Probable Succession of Fo rmations- Figure 14 Successional arrangement of pine-barren formations. trees, which, when admixed with pitch-pines, can be designated the Mixed Pine-Oak Formation. The pines may ultimately suppress the oaks, and the vegetation returns to the original pine forest in which the series is bound sooner or later to culminate. The full details of this succession and the interrelation of the different formations of the region will be emphasized as we proceed. Pine-Barren Formation Historically, the occupancy of the recently elevated coastal plain during interglacial times by the pitch-pine and other associated species produced what we now call the pine-barrens. Originally more widely distributed, 4 50 VEGETATION OF THE NEW JERSEY PINE-BARRENS this type of vegetation became restricted in area with the Pensauken submergence. The island, then formed, was probably covered with pine forest, and when a reelevation of the land took place, the boundaries of the pine-barrens, which had been determined by the coast-line of Pen- sauken island, were maintained by the encroachment of a deciduous forest type of vegetation on the land surface elevated recently above the sea in post-Glacial times. The pine which makes up the bulk of the New Jersey pine forest is Pinus rigida Mill, the pitch-pine, associated in places with the yellow Figure 15 Open pine forest in Lebanon Reserve, with snow on the ground, December 29, 1908. The absence of a dense undergrowth is here noteworthy. (Courtesy of N. J. Forestry Commission.) pine, P. echinata. If the ascent of a hill, or other elevation, is made, an unbroken extent of dark-green pine forest is seen stretching off into the distance as far as the eye can reach. A closer inspection, however, shows considerable diversity of vegetation as one passes through the open, sunlit pine forest from one point to another (Fig. 15). The average height of the pines in old stands from 3 to 4 decimeters (10 to 14 inches) in diameter was 20 to 22 meters (65 to 70 feet). Where fire has de- stroyed the original forest, the pine trees are much shorter, 7 to 14 VEGETATION OF THE NEW JERSEY PINE-BARRENS 5' meters (20 to 40 feet) being an average size for such trees, some of which, if badly injured by fire, show Hght-green sprouts growing from the trunk or base of the trees (Figs. 16 and 17). The forest canopy consists of the dense crown of the standing pine trees, but only in exceptional cases do the crowns meet to shade the ground underneath. The trees are spaced so that considerable sunlight reaches the forest floor, and hence the forest may be described as an open one (Fig. 18). The pine trees are light I'loURE 16 Regeneration of fire-denuded pitch-pines, Pinus rigida, near the Sym Place, July 29, 1913. Note the green sprouts on the central tree and the bushy young trees that have sprouted from the roots of old trees whose tops were killed by fire. demanding, and they are dominant. All other growth has been kept suppressed by the dominant pines. The density of an old pine forest near Winslow is indicated in the following summary of measurement of twelve plots of one acre each.* Likewise the quadrat method gives accurately the number of trees in the forest per 10 square meters, or any other unit taken arbitrarily as the unit of measurement. Such an estimate will be shown graphically on a later page. Report of the N. j. State Geologist, iJ Report on Forests, p. 115. Figure 17 Pitch-pine, Pinus rigida, in recently burned-over area, near the Sym Place, west of the Lower Plain, July 29, 1913. Trunks of the taller fire-denuded trees show the characteris- tic green sprouts formed by a regeneration of the branches. (Photo by George E. Nichols.) ^mx^^^t^ Figure 18 Young open pine forest at Hanover, October 8, 1909. The low scrub oaks and other low shrubs are typic of many stretches of the high pine-barren formation. (Courtesy of N. J. Forest Commission.) 52 VEGETATION OF THE NEW JERSEY PINE-BARRENS 53 Mer- Num- Maxi- Total chant- Num- Average ber OF Average mum Volume able Plat Num- ber OF Diam- Trees Diam- Diam- Average Over 6 Volume Num- ber OF Trees eter, Over eter, eter, Height Inches Over 10 ber Trees Over 6 Breast 10 B reast Breast IN CU5IC Inches Inches High Inches High High Feet in Board Feet I 67 66 13.9 34 14.4 22 69.1 2,882 10,170 2 70 65 14.0 5> 16.2 18 6S.8 2,917 9-925 3 4 78 72 13.0 56 14.8 18 66.9 2,812 8,940 86 74 12.7 56 13.9 20 69.7 2,548 8,828 6 92 78 12.0 55 13.4 18 68.1 2,557 7.739 7 8 65 60 13.0 46 14.3 21 67.5 2,228 7.521 71 69 12.4 54 13-5 19 66.3 2,388 7.488 9 10 68 63 12.5 49 13. 1 19 68.1 2,127 7,2oS 104 78 I 1.2 5" 12.6 ■7 68.1 2,048 6,134 12 45 43 13. 1 35 14. 1 20 68.1 1,613 5.631 '4 I? 85 62 I I.O 35 13-1 "9 66.8 1,708 4,618 74 57 10.4 27 12.8 16 66.8 1.275 3.369 Architectural Form of Pine Trees.— The shape of the forest- grown pine trees varies considerably, as the accompanying figures will show. The variation of the branching system of native forest trees has been studied very little in this country, although abroad considerable attention has been given to it. Miss Annie Oakes Huntington, by a series of descriptions and photographs, accomplishes this in her " Studies of Trees in Winter," published in 1902. Jepson, in his " Silva of Califor- nia," describes the architectural form of many Pacific Coast trees, as do Blakeslee and Jarvis in their "New England Trees in Winter" (1911). Dr. Ludwig Klein has undertaken, in Karsten and Schenck's "Die Vegetationsbilder," * to describe the Charakterbilder mittel europaischer Waldbaume, and has collected many beautiful photographs which accompany the text. Professor G. Haberlandt has characterized the form of tropic trees in Java in his " Eine botanische Tropenreise," published in 1893. "The Artistic Anatomy of Trees, their Structure and Treatment in Painting," by Rex Vicat Cole (191 5). is a mine of information on this subject. The pitch-pine, when growing in dense stands in the forest and under stress of competition for light, develops typically a central axis, with all the branches disposed laterally to it, with the lower branches killed by shading and in many cases broken off, leaving conspicuous stubs, which bristle in whorls beneath the green crown of the tree (Figs. 19, 20, and 21). The axis is not always perfectly straight, but may be inclined or bent, the bending being more or less ess-shaped (Figs. 22, 29, and 30). Other trees are conic (Figs. 19, 20, 21, and 22). The pitch-pine responds to the 54 VEGETATION OF THE NEW JERSEY PINE-BARRENS advantage of free space and branches more freely until it becomes round- headed (Fig. 28). Frequently the stem bifurcates. This bifurcation may be found at the base of the trees (Fig. 23), half way up (Fig. 24), or near the top (Fig. 25). Such trees are V-shaped or Y-shaped (Figs. 24 and 25). Another common form of bifurcated stem is the stag- /, -ivi^^t; Figure 19 Conic form of pitch-pine. Figure 20 Conic form of pitch-pine. headed, where the main leader divides near the top into several branches, each of which branches again repeatedly. Occasionally a tree becomes candelabra-shaped (Figs. 26 and 27), when it represents a tree which has developed in an open space in the woods. Some of the forest patriarchs are of this form. At those localities where the pine-barrens approach the VEGETATION OF THE NEW JERSEY PINE-BARRENS 55 sea, as at Spring Lake, New Jersey, or the salt marshes, as at Barnegat Pier and Somers Point, the trees become reduced in size; the dead branches and old cones are present in some trees which become irregular and their architectural shapes are wind controlled. Here they become wind-swept and one-sided (Figs. 31 and 32). One of the usual forms is the bisected tree, where one side fails to develop, owing to the action of Figure 22 Round-head form of pitch-pine. Figure 21 Conic form of pitch-pine. Figure 23 Bifurcated form of pitch-pine. the wind on the exposed side of the tree (Figs. 33 and 34). Another form is the bush or shrub form (Garben-baum, or sheaf tree) which produces freely a number of clustered stems and more or less wind-blown (Fig. 35). This form may be called the bush shape. Another form may be called the candelabrum type, as the branches of the low shrubby form are dis- posed in a way to suggest a candelabra (Fig. 36). At the edge of the pine 56 VEGETATION OF THE NEW JERSEY PINE-BARRENS forest near the sea, low, prostrate, wind-swept trees are found, their top branches incHned, arched, or bowed in a direction contrary to that of the prevaiHng winds (Fig. 37). Such trees on a high mountain form the " Krummholz," or elfmwood of the ecologist. At the seashore these trees are called appropriately v/ind-swept dwarfs. One other architec- tural form is seen on the plains of New Jersey, where the pine trees are all low, but cone-bearing. Such low trees, showing nanism, may be denominated, because of their shape, cushion trees. Tabulated, the forms of pine-barren trees are as follows: Conic (Figs. 19, 20, 21, 32) Round-headed (Figs. 22, 28, and 38) Bifurcated (Figs. 23, 24, 25, 26, 27, 34) V-shaped (Fig. 23) Y-shaped (Figs. 24 and 25) Stag-headed (Fig. 26) Bisected (Figs. 33 and 34) Cushion-shaped (Fig. 133) Candelabra-shaped (Fig. 36) Bush- or sheaf-form (Fig. 35) Wind-swept dwarf (Fig. 37) Figure 24 bifurcated and Y-shaped pitch-pine. Figure 25 Figure 26 Bifurcated and Y-shaped Bifurcated form of pitch- pitch-pine, pine. Finally, it might be stated that, although the typic pitch-pine trees have excurrent stems, yet in the forest we frequently fmd trees that are deli- quescent, and an outline drawing of the branch system of such trees without foliage might be taken for that of a deciduous tree. In other trees the branches are drooping, but they never become typically weep- ing varieties, because the branches are thick, gnarled, and not pliant enough to assume the pendent habit. Facies of Pine-Barren Formation.— Several facies of the pine- barren formation can be distinguished. The character of these facies VEGETATION OF THE NEW JERSEY PINE-BARRENS 57 depends on the trees and shrubs which are associated in the pitch-pine forest. The species that are associated with the pitch-pine are deter- mined to some extent by the soil conditions. Thus we can distinguish wet pine-barrens, flat pine-barrens, dry pine-barrens, and high pine- barrens. There are pine-barren facies with a bear-oak (Quercus ihci- foha) undergrowth, with a black-jack oak (O. marylandica) undergrowth, and with a laurel (Kalmia latifolia) undergrowth. A common facies is where the heath-like plants (Dendrium, Vaccinium, etc.) are abundant, and there are facies where there is little undergrowth, but bare sand with prostrate trailing species, or reindeer lichen with associated mosses. There are pine-barrens transitional to the open formations of the sea dunes, to the cedar swamps of the interior, to the deciduous swamps of the sluggish streams. In fact, it can be said that a careful examina- -^W^M^-r-^^ tion of the pine-barren region reveals almost Figure 27 Bifurcated pitch-pine. Figure 28 Round-headed pitch-pine. Figure 29 Excurrent pitch-pine tree. as great a variety of types as there are physiographic and soil condi- tions, and as there are species which enter into competitive or com- plementary association in the facies. Before commencing the syneco- logic description of the pine-barren vegetation, it may be well to detail the method employed in the study of this vegetation, as such methods may be found helpful to other botanists working in the same field. Method of Investigation. — The observations of the writer were recorded in a series of field note-books. These note-books are of pocket size, 1 1.8 cm. (4^4 inches) wide by 18.7 cm. (y^o inches) long, with flex- ible backs, opening at the top. Each book, which cost five cents, con- tained approximately 60 pages of ruled paper. A map of the portion of 58 VEGETATION OF THE NEW JERSEY PINE-BARRENS the pine-barren region chosen for study was drawn in reduced size on the page of the note-books preceding the field notes for that district. Blue pencil indicated roads and railroads, the latter marked by cross lines. Red pencil showed rivers, streams, and swamps, while green indi- cated forest areas. The geographic names were written in black ink, and the localities where photographs and habitat observations were made were entered in lead pencil on the face of the map, while in the field. In journeying across the country by railroad, observations were made from the car windows and entered in the books with the names of the nearest stations or mile- posts as geographic reference points. Thus observations made in walking from place to place were connected by obser- vations made from the moving trains. The routes taken in this sort of botanic survey :#Xi-t-f Figure 30 Pitch-pine with ess-shaped top. Figure 31 One-sided pitch-pine tree. work were indicated on a reference map. The following system was adopted to make the field notes as accurate as possible. Quite a number of plants, such as Pinus rigida, Quercus ilicifolia, Kalmia latifolia, K. angustifolia, Dendrium buxifolium, Xerophyllum aspho- deloides, about the identity of which there could be no question, were entered in the note-books without verification. Frequently, when in doubt, the plant was identified in the field by consulting Britton's or Gray's Manual without preservation of the specimens. If unable to VEGETATION OF THE NEW JERSEY PINE-BARRENS 59 determine the name satisfactorily by the manual, a specimen was gath- ered for more careful study with herbarium material at hand. Hence, while not following the method of some botanists in insisting that each record should be accompanied by a dried specimen of the plant, yet the above system of verification can be used with a large degree of accuracy. The method advocated by Stone and others to have a herbarium sheet of the plant is desirable where purely syste- FlGURE 32 Wind-swept pitch-pine tree. Figure 33 Bisected pitch-pine tree. Figure 34 Bisected pitch-pine tree. Figure 35 Bush form of pitch-pine. matic work is concerned, but the method used by the writer is the only feasible one where a synecologic study of the vegetation of a region of considerable size has been undertaken. 6o VEGETATION OF THE NEW JERSEY PINE-BARRENS High Pine-Barren Facies. — In the heart of the New Jersey pine- barrens, as at Lakehurst, Whitings, and Bamber, are rolling hills not Figure 36 Candelabra-shaped pitch-pine tree. Figure 37 Wind-swept dwarf pitch-pine. dissected by streams, which are separated from each other by large inter- vals of elevated country, some- times with gravelly, sometimes with sandy, soil. The soil, being as a rule dry and well drained by seepage and surface run-off, is covered by a facies of the pine forest in which only the plants of the drier soils are associated with the pines (Fig. 39). The plants of moister situations are absent entirely. The prevailing pine, Pinus rigida, forms an even stand where the trees range in height from 7 to 12 meters (25 to 40 feet), as seen in Fig. 41. The canopy of this forest is fairly close, although there are inter- vals where the pine trees are spaced more widely (Figs. 39 and 40). From Quadrat 2, taken one mile east of Lakehurst, it has been ascertained that there are I 5 pitch-pine trees to a quadrat of ten square meters, as com- pared with 14 in Quadrat i of the same size surveyed near the sea-coast Figure 38 Pitch-pine tree, Pious rigida, with dense witches'-broom, at Clementon, N. J. (Photo- graph by Genji Nakahara and J. W. Harsh- berger.) VEGETATION OF THE NEW JERSEY PINE-BARRENS 6l at Como Lake, or as contrasted with Quadrat 3, taken at Sumner, N. J., with 13 trees. Associated with the pitch-pine occur the short-leaf pine. Figure 39 Level dry pine-barrens about two miles southeast of Lakehurst, N. J., 10 a. m., August 2, 1909. (Photo by Roland M. Harper.) Figure 40 Dry pine-barrens about a mile east of Toms River, N. J., 3.23 p. m., August 2, 1909. (Stereoscopic photographs by Roland M. Harper.) Pinus echinata, as at Lakehurst and Shamong, and an under story, or secondary layer, of medium sized trees, composed of Quercus marylandica (Figs. 42^ 43. and 44), Q. stellata (Fig. 45), Q. velutina (Fig. 46), Sassa- 62 VEGETATION OF THE NEW JERSEY PINE-BARRENS f ras variifolium sprinkled as tolerant species beneath and between the pine trees. The third layer, or story, of this type of forest consists of the bear- oak, Quercus ilicifolia (Fig. 47), and the dwarf chestnut-oak, Quercus prin- oides (Figs. 48 and 49). These are low, rounded bushes, or small trees, with a number of branches rising from a powerful tap-root. The bear- oak, Quercus ilicifolia (Fig. 47), is the more common of the low oaks in New Jersey. It bears usually an abundant crop of acorns, and is very resistant to the destructive action of fire, as it readily sprouts from the charred stump after a destructive conflagration. The dwarf chestnut- FlGURE 41 Road through a scattered stand of pitch-pine, Pinus rigida, in Lebanon Reserve, De- cember 29, 1908, with snow on the ground. (Courtesy of N. J. Forest Commission.) oak, Quercus prinoides, disputes the ground with the bear-oak, especially in the high pine-barrens. It is usually of smaller size and more spreading in habit. Occasionally these oaks close together to form a low thicket from I to 1.2 meters (3 to 4 feet) high beneath the pines. As a fourth story, or layer, we find associated the sweet fern, Comp- tonia asplenifolia (Fig. 50), the huckleberries, Gaylussacia dumosa, G. resinosa. Ilex glabra, Kalmia angustifolia (Fig. 51), Rhus glabra (Lake- hurst), while the fifth layer, or story, of low shrubs consists of the blue- berries, Vaccinium pennsylvanicum, V. vacillans, and such grasses and VEGETATION OF THE NEW JERSEY PINE-BARRENS 63 Figure 42 Low black-jack oak, Quercus marylandica, in pine forest at Shamong, June 27, 19 10. (Photograph by Henry Troth and J. W. Harshberger.) Figure 43 Pine-barrens at Shamong (Chatsworth), Oc- tober II, 1913. On the right of Prof. Adolf Engler, of Berlin, Germany, who is holding Sarracenia purpurea in one hand, is Quercus ilicifolia (nana), with white cedar in the back- ground. Immediately back of Dr. Engler and to the left is Quercus marylandica. Fi(.1:ki-, 44 Plowed fire line with Prof. C. von Tubeuf, of Munich, Germany, on its inner, undis- turbed edge, with Quercus marylandica, and the outer area cleared of undergrowth with a few dwarf oaks. October 11, 1913. P^fr E_*^^^. .^■^^'^^ ..*j»^ %'*' simg'^^^T?riB^^'^'*Si>i^ i^^ ' w. ^|HiB^^^iM^SjS,tj^ 3"^*^ ^^ ■3' •> ^^BB^^^f '"^'^'-3 M ^^^B ^^^^^^^^ Kalmia latifolia, K. angustifolia, Dendrium buxifolium, Arcto- staphylos uva-ursi, and Gaul- theria procumbens. These all grow in different relationship to each other and in different shades of green. Against this dark-green background one sees splashes of bright-red color which prove to be the bright red leaves of Smilax glauca and Gaylussacia resinosa and a few lower leaves of Quercus stellata, while in slightly wet places the bright red to reddish-purple leaves of Vaccinium atrococcum are seen. But the color tones of the oaks in late autumn are deep brown to yellow-brown of the persistent leaves. In a piece of pine woods used as a picnic ground, where the ground has been cleared repeatedly, and where it had been tramped hard, the bright colors of the low leafy sprouts contrasted strongly with the lifeless browns of the same species in the nearby forest. The sprout leaves of Quercus prinoides were orange-yellow to yellow-brown, then red to claret- red; those of Q. stellata, and Q. velutina were bright red; those of Q. ilicifolia, red to reddish- claret, reddish-brown to orange- red. Figure 66 Blue lupine, Lupinus perennis, in full flower at Browns Mills, May 13, 191 1. (Photograph by Miss Lydia P. Borden.) Figure 67 Pine forest of pitch-pine, Pinus rigida, at Mays Landing, April 6, 19 10. Lower branches of pines draped with green brier, Smilax rotun- difolia. Litter of pine needles broken with .green mosses and beard-grass, Andropogon scoparius. in the background holly, ilex opaca, post-oak, Quercus stellata, bayberry, Myrica carolinensis, etc. (Photograph by Henry Troth and J. W. Harshberger.) lantic coast between Sea Girt on the south and Asbury Park (Deal Lake) on the north, at Sea Girt, Spring Lake, Belmar, Avon, Bradley Beach, Ocean Grove and Asbury Park. Coastal Pine-Barrens The pine-barrens reach the At- VEGETATION OF THE NEW JERSEY PINE-BARRENS 83 Rapid inroads have been made in this coastal forest since about 1876 by the estabHshment of various summer resorts, so that the original condi- tion of the vegetation must be inferred from the remnants. At several points, notably Sea Girt, Spring Lake, and Belmar, this forest approaches the sea in almost unaltered state. It is, therefore, important to place on permanent record the facts obtained by a study of these original condi- tions. This study was made more complete by noting the more con- spicuous plants on each of the building lots in the borough of Belmar, and by noting the vegetation left in isolation at various points between Belmar and Asbury Park. At Avon, along the north shore of Shark River Bay, were noted Pinus rigida, Quercus alba, Q. coccinea, Q. mary- landica (Fig. 68), Q. rubra, Q. stellata, Prunus serotina. Sassafras varii- folium, the waxberry shrub, Myrica carolinensis, and the tall herb, Baptisia tinctoria. In addition to the above the black-oak, Quercus velutina, was noted at Bradley Beach, Deal Lake, which is approxi- mately the northeastern limit of the pine-barrens in New Jersey. Here were noted, August 7, 1909, Acer rubrum, Liquidambar styraciflua, Nyssa sylvatica, Pinus rigida, Quercus alba, Q. phellos, Q. rubra, Q. velutina, and Sassafras variifolium. The remnant of the pine-barren vegetation, as determined by a lot to lot census at Belmar, consists of the trees and plants enumerated in the following list: Herbs Chrysopsis mariana. Gerardia purpurea. Lespedeza capitata. Melampyrum americanum. Pteris aquilina. Solidago odora. LlANES Ampelopsis quinquefolia. Smilax glauca. Smilax rotundifolia. Vitis labrusca. Epigaea repens. Rhus radicans. Low Woody Perennials Rubus cuneifolius. Vaccinium macrocarpon. Shrubs Amelanchier canadensis. Azalea viscosa. Clethra alnifolia. Gaylussacia frondosa. Gaylussacia resinosa. Ilex glabra. Myrica carolinensis. Prunus maritima. Pyrus arbutifolia. Quercus ilicifolia. Rosa lucida. Rhus copallina. Sambucus canadensis. Sassafras variifolium. Vaccinium corymbosum. Vaccinium pennsylvanicum. Vaccinium vacillans. Viburnum dentatum. 84 VEGETATION OF THE NEW JERSEY PINE-BARRENS Figure 68 Barren oak, Quercus marylandica, growing beneath pitch-pine near Belmar, N. J. in coastal pine-barrens. (Photograph by E. L. Mix and J. W. Harshberger.) VEGETATION OF THE NEW JERSEY PINE-BARRENS 85 Trees Acer rubrum. Carya alba. Diospyros virginiana. Juniperus virginiana. Liquidambar styraciflua. Magnolia glauca. Nyssa sylvatica. Pinus rigida. Prunus serotina. Quercus alba. Quercus coccinea. Quercus marylandica. Quercus phellos. Quercus rubra. Quercus stellata. Quercus velutina. Contrast of German and New Jersey Formations it is important in phytogeographic work to contrast the different types of vegetation of the world with one another. One of the most im- portant results of the interna- tional phytogeographic excur- sions, begun auspiciously in Swit- zerland and in England and con- tinued by American ecologists through the United States in the summer of 19 13, has been the comparison of the different kinds of vegetation types found on the different continents. Pro- fessor C. Schroter, on the trip of the phytogeographers to the pine-barrens of New Jersey, suggested the similarity of the physiognomy of the pine-barrens and the pine-heaths (Kiefern- heide) of Europe. In following out this suggestion, a reading of Graeb- ner's " Die Heide Norddeutschlands" has enabled the writer, as one of the leaders of the excursion in New Jersey, to make a more intelligent com- parison of the two types of vegetation. The heathland of North Ger- many is developed in a region controlled by an oceanic climate. Where the continental climate begins to prevail pine trees invade the heath- land and it is converted into a pine-heath, with the pines as dominant plants and the heath flora as subordinate. A sojourn on the Island of Nantucket, 45 kilometers (28 miles) out at sea, during two summers, sug- gested in a study of the vegetation there an hypothesis as to the origin of the New Jersey pine-barrens. Remove the pine trees from the forests of New Jersey and some of the pine-barren species found there and you would have the conditions on Nantucket controlled by an oceanic climate (Fig. I 50). The Nantucket vegetation is a true heath with many of the Figure 69 Sandwort, Arenaria caroliniana, associated with spurge, Euphorbia ipecacuanhae, sweet- fern, Comptonia asplenifolia, blueberry, Vaccinium pennsyivanicum, etc., at Shamong (Chatsworth), June 27, 1910. (Photograpli by Henry Troth and J. W. Harshberger.) 86 VEGETATION OF THE NEW JERSEY PINE-BARRENS southern coastal plain species missing. Add the pitch-pine, which is now invading the central part of the Island of Nantucket, and you would have a typic pine-barren vegetation or pine-heath (Kiefernheide) (Fig. iji)- Similarly, as in Germany, with the control of a continental climate, the pine trees in the New Jersey region become dominant and the heath- plants, in the form of Arctostaphylos, Dendrium, Gaylussacia, Kalmia, Vaccinium, and dwarf forms of Quercus become subordinate to the pines and form the characteristic undergrowth which has been described in detail on former pages. The different facies of pine-barren vegetation which have been described may be compared with the Waldheiden of the Germans, for we find that Graebner distinguishes two types: " Kiefern- heide" and "Laubwaldheiden," and the distinctive facies of these two types are classified by him as follows: Waldheiden 1 . Typus Kiefernheide. Facies b. Kiefernheide mit Vorherrschen von Juniperus com- munis. Facies c. Kiefernheide mit Vorherrschen von Rubus-arten. *Facies d. Kiefernheide mit Vorherrschen von Arctostaphylos. *Facies e. Kiefernheide mit Vorherrschen von Grasern. *Facies f. Feuchtige moosige Kiefernheide. *Facies g. Kiefernheide mit Vorherrschen von Vaccinium myrtillus und V. Vitis-idaea. 2. Typus Lauhwaldheiden. Facies a. Birkenheide. *Facies b. Eichenheide. Those facies of the forest-heath in Germany which are similar physi- ognomically to the ones in New Jersey are marked with an asterisk. Facies g in Germany, with the prevalence of two species of Vaccinium, is represented in New Jersey by a pine forest with an undergrowth of Gaylussacia resinosa, Vaccinium pennsylvanicum, V. vacillans, and Kalmia angustifolia. The oak-heath (Eichenheide) is represented in New Jersey by the secondary formation which we have designated oak- coppice, which in some places is prominent after the pines have been re- moved. Oak-heath in New Jersey succeeds pine-heath. The bear- berry, Arctostaphylos uva-ursi, is one of the most abundant plants on the heaths of Nantucket, where it forms carpets an hectare (i to 2 acres) in extent, representing the bearberry heath. It also occurs as a common undergrowth in the high pine-barrens of New Jersey, which thus may be contrasted with Facies d of the German pine-heath. A more detailed study by instrumental methods by travel in this country and abroad would be extremely profitable in working out the details of vegetational types. CHAPTER VI PINE-BARRENS TRANSITIONAL TO SEA-DUNE VEGETATION From the ocean end of Como Lake, inland from the built-up portion of Belmar south to Como Lake, the prevailing vegetation, as indicated by the sky-line, is coniferous. The ocean front of this forest consists of low pine trees, as at Belmar Park. South of Como Lake, on the Como side, the prevailing tree is Pinus rigida, and the composition of the forest may be studied now at close range. Crossing the dune complex covered with Ammophila aren- aria, Myrica carolinensis. Soli- dago sempervirens, Stropho- styles helvola, Triodia cuprea, we come to the strip of forest facing the northeast, with dead, wind-swept cedars and clumps of spire-shaped, living ones (Fig. 70), with an association of wind- swept trees of Pinus rigida. Mag- nolia glauca, Nyssa sylvatica, and Sassafras variifolium, inter- mingled with such shrubs as Cephalanthus occidentalis. Ilex opaca, Myrica carolinensis, Rhus copallina, and Vaccinium corym- bosum, together with the switch- grass, Panicum virgatum, and the boneset, Eupatorium album (com- mon). The vines are Ampelopsis quinquefolia, Lonicera japonica (in- troduced and naturalized), Rhus radicans, Smilax rotundifolia and Vitis labrusca. The vegetation of the dune complex merges with that of the thicket formation. We now enter the pine-barren formation, which extends inland. The composition of this forest in Spring Lake (Como) is as follows: the forest floor, covered with pine needles, is dotted over with clumps of Ammophila arenaria, Baptisia tinctoria, with a sprinkling 87 Figure 70 Extreme outer edge of pine-barrens fronting the sea dunes at Spring Lake, August 24, 19 10. Note wind-swept trees of Nyssa sylvatica, Juniperus virginiana, Pinus rigida, with which as lianes are associated Rhus radicans, Vitis labrusca, Ampelopsis quinquefolia. (Photo- graph by E. L. Mix and J. W. Harshberger.) VEGETATION OF THE NEW JERSEY PINE-BARRENS of Sericocarpus asteroides. There are isolated bushes of Myrica caro- linensis and Quercus ihcifoha. As we walk inland we find that the marram-grass, Ammophila arenaria, disappears, and out of the pine needles grow Baptisia tinctoria, Cypripedium acaule, Melampyrum americanum, Smilax glauca, and low trees of Prunus serotina. Here grow also Andropogon scoparius, Chrysopsis mariana, Gaylussacia resinosa, and Quercus ilicifolia. Back of the transitional pine forest (Fig. 71), which immediately faces the ocean on the Spring Lake and Belmar sides of Como Lake, we enter the pine-barrens proper (Fig. 72), where Pinus rigida forms erf *-^^ -•.•|„ w,;,, '^< rvin 2- J. -'':' ■§'■ ■'• v^lWm ' im I iiiiii'y 1 m^nm^ ■ 1 Figure 71 Open coastal pine forest with little under- growth near Manasquan. an open growth in association with the oaks Quercus alba, Q. mary- landica (Fig. 73), Q. phellos, Q. stellata, Q. velutina, and the red cedar, Juniperus virginiana, oc- casional and of secondary habit. Other trees of less importance are Populus grandidentata, Prunus serotina, and Sassafras variifo- lium. The shrubs of this forest formation are Amelanchier inter- FlGURE 72 Pine forest (coastal) at Belmar, August 24, 1910. Clump of waxberry, Myrica carolin- ensis, in foreground, and bear-oak, Quercus ilicifolia, at the right edge of the picture. (Photograph by E. L. Mix and J. W. Harsh- berger.) VEGETATION OF THE NEW JERSEY PINE-BARRENS 89 media, Andromeda mariana, Clethra alnifolia, Comptonia asplenifolia, Gaylussacia resinosa. Ilex glabra, Kalmia angustifolia, Myrica caro- linensis, Prunus maritima, Quercus ilicifolia, and Rhus copallina, together with the lianes, Rhus radicans, Smilax glauca, and S. rotundifolia. The herbaceous plants noted in this coastal forest were the grasses Am- mophila arenaria, Andropogon fur- FlGURE 73 Black-jack oak, Quercus marylandica, at edge of sand dune, Belmar, N. J. Note that the lower part of the tree to the left has a cluster of adventitious roots formed when the sand covered the base of the tree. These roots no longer function, as the sand has been removed by wind action. (Photo- graph by E. L. Mix and J . W. Harshberger.) Wi 0 ^P^ !4-.'^^9^H^^I ! m ^""^Wm rf W'- ^^ f$(i Sfrnf 1 Kyi iffi Wm 1 9Sth.^\ ' -^KiHi ■ ' luj^^^^B pu mt£. F*^'''2-^^i^ Wt W^'WBi ■ ^liH^pi i ^^ft 1 f^^ 1 fr^'^^i^ 1 '•■-^ift*^****?^-. ^^^SB^^^^m I "*. 1 Figure 74 Coastal pine forest at Belmar. This forest is second growth on an old abandoned field, as it is almost entirely devoid of undergrowth, and shows old furrows, as indicated by the parallel shadows run- ning back in the picture. (Photograph by Mrs. Helen B. Harshberger.) catus, together with Ascyrum hypericoides (in clumps), Asclepias obtusi- folia, Aster ericoides, Baptisia tinctoria, Chimaphila maculata, Chrysop- sis mariana, Cypripedium acaule, Erigeron canadensis, the prostrate, sand-loving Euphorbia ipecacuanhae, Gaultheria procumbens, Gerardia purpurea, Hudsonia tomentosa, Melampyrum americanum, Polygonella articulata, Pteris aquilina, Sericocarpus asteroides, and Solidago odora (Fig. 74)- 90 VEGETATION OF THE NEW JERSEY PINE-BARRENS Transition from Sea Dunes to Pine Thicket at Sea Girt Starting inside the narrow dune area, the thicket formation is approached by crossing a narrow tension strip of Ilex glabra, I. opaca, low red cedars, Juniperus virginiana, bayberries, Myrica carolinensis, Prunus serotina, Rhus copallina, and the lianes,Ampelopsis quinquefolia and R. radicans. Here are found such herbs as Chrysopsis mariana, Eupatorium album, and Solidago sempervirens. These plants enter the thicket and mingle with the wind-swept cedars, which are draped with the vines Ampelopsis Figure 75 Blending of bay beach vegetation with Panicum virgatum, golden-rod, Solidago sem- pervirens, and dense forest. On bluff and inside forest grow 1 lex opaca, red cedar, Juni- perus virginiana, Quercus stellata, etc. Somers Point, April 6, 1910. (Photograph by Henry Troth and J. W. Harshberger.) quinquefolia, Celastrus scandens, Rhus radicans, Smilax rotundi- folia, and Vitis labrusca. The pitch-pine, Pinus rigida, the black- jack oak, Quercus marylandica, the holly, Ilex opaca. Sassafras varii- folium, sour-gum, Nyssa sylvatica (occasional), are found as associates of the red-cedar, together with the huckleberry, Gaylussacia frondosa, bayberry, Myrica carolinensis, Rosa Carolina, Viburnum dentatum, with such herbaceous plants as Chrysopsis mariana, Gerardia purpurea, Lespedeza capitata, and Meibomia obtusa. The other constituents mentioned above remain about the same. VEGETATION OF THE NEW JERSEY PINE-BARRENS 91 Bluff Forest Along Great Egg Harbor Bay at Somers Point Below Somers Point the pine forest comes down to a steep bluff. Here, on the front slope, April 2, 1910, grew Pinus rigida, *Juniperus virginiana, Quercusstellata,and*Myricacarolinensis (Fig. 75). The narrow, shingly beach at the foot of the bluff (Fig. 75) was tenanted by the switch-grass, Panicum virgatum, Baccharis halimifolia, and the oak, Quercus stellata, with gnarled and twisted branches, and the holly, *llex opaca, in the order of their occurrence. On the level ground back from the edge of the bluff (Fig. 76) the components of the forest were, in theorderof their importance: Pinus rigida, *llex opaca, *Juniperus vir- giniana, Quercus stellata, O. mary- landica, Q. alba, *Cornus florida, Hicoria glabra, with the bushy ^'' '^^'"^Hi ^HHffi^^^ HNP^^^^^^ ^Si" •• -~^m ggK^K BflHHl m: "^^ ^ ^''^^^^j^^S^hHHHrII^h Figure 76 Interior view of bluff woods, Somers Point, with pitch-pine, Pinus rigida, holly, Ilex opaca, red cedar, Juniperus virginiana. April 6, 19 10. (Photograph by Henry Troth and J. W. Harshberger.) Figure 77 Interior of pine woods south of salt marsh, Somers Point, April 6, 1910, with Pinus rigida, Ilex opaca, Quercus velutina, Quer- cus alba, etc., and oak and pine litter. (Photograph by Henry Troth and J. W. Harshberger.) *Myrica carolinensis and the Hanes, *Smilax rotundifolia and Vitis sp. This type of forest (Fig. 77) graded imperceptibly into a facies where pitch- pine, Pinus rigida (some trees 6 decimeters in diameter), was dominant (Fig. 78). The secondary trees were Ilex opaca, *Juniperus virginiana, Quercus alba, Q. marylandica, Q. stellata, and Q. velutina. The forest litter consisted mainly of the fallen leaves of the oaks and the pines (Fig. 79). The shrubby undergrowth consisted of *Gaylussacia resinosa, 92 VEGETATION OF THE NEW JERSEY PINE-BARRENS '%^*^' ,#^-^'^^ k^^''?^ y*^ 't^ > l'4*^Sf:''r Figure 78 Interior view of pine forest witii large pine, Pinus rigida, 7.62 dm. {2}4 feet) in diameter, surrounded by a thicket of Ilex opaca, Quercus alba, and wintergreen, Gaultheria procumbens, with a litter of pine needles and oak leaves, Somers Point, April 6, 1910. (Photograph by Henry Troth and J. W. Harshberger.) VEGETATION OF THE NEW JERSEY PINE-BARRENS 93 *Ilex glabra, *Myrica carolinensis, *Rhus copallina, and young trees of *Ilex opaca (Fig. 80), Pinus rigida, Quercus alba, Q. marylandica, and Q. stellata. All these trees were festooned with climbing vines, which add to the jungle-like growth. The lianes noted in this forest were: *Ampelopsis quinquefolia, *Rhus radicans, *Smilax rotundifolia, and *Vitis labrusca. This dense thicket has been the favorite nesting place Figure 79 Open mixed forest at Somers Point, with large white-oak, Quercus alba, Pinus rigida, Ilex opaca, Juniperus virginiana, Smilax herbacea, Vaccinium pennsylvanicum, etc. (Photograph by Henry Troth and J. W. Harshberger.) Figure 80 Interior of pine forest, Somers Point, April 6, 1910, with Pinus rigida. Ilex opaca. Sas- safras variifoiium, Quercus prinoides, and Smilax rotundifolia. (Photograph by Henry Troth and J. W. Harshberger.) of shore and migratory birds during the summer, and in winter, owing to the shelter afforded by the evergreen trees, many birds linger on their way south, especially if the winter is an open and a mild one. The trees, shrubs, and lianes of this thicket have been bird distributed with the exception of the pines. The same type of forest thicket is char- acteristic of the sea islands along the coast as far south as southern Florida. Its character is largely due to the trees with avivectent fruits and seeds.* *To emphasize this important fact, all plants with bird-carried fruits have been marked in the above account by an asterisk. 94 VEGETATION OF THE NEW JERSEY PINE-BARRENS Tension Line Salt Marsh to Pine Forest at Somers Point A sharp tension line was found at Somers Point, between the pine forest and the salt marsh (Fig. 8i). The salt marsh plants meet a narrow strip, in some places 6 meters (20 feet) wide, in which Panicum virga- tum, Solidago sempervirens, and Baccharis halimifolia were in associa- tion. Back of this grassy strip the outer edge of the forest consisted of Juniperus virginiana, advancing upon the grassy strip, together with Figure 81 Point of pine forest projecting into salt marsh. Pinus rigida dominant, associated with Ilex opaca, Juniperus virginiana, Myrica caroiinensis, fronted by a strip of Panicum virgatum, Somers Point, April 6, 1910. (Photograph by Henry Troth and J. W. Harsh- berger.) Ilex opaca, Pinus rigida, Quercus alba (large), Q. stellata, with Myrica caroiinensis, Smilax rotundifolia, and the ground herbs Chimaphila maculata and Mitchella repens (Fig. 81). The forest proper on the north side of this tongue of salt marsh consisted of the codominant Ilex opaca, Juniperus virginiana, Pinus rigida, Quercus alba (Figs. 82 and 83), while the secondary and younger trees growing in suppression to the taller trees were Ilex opaca, Quercus alba, Q. marylandica, Q. stellata, Q. velutina, and Sassafras variifolium. The forest litter consisted of VEGETATION OF THE NEW JERSEY PINE-BARRENS 95 oak and pine leaves, about four centimeters deep, out of which grew Chimaphila maculata and Gaultheria procumbens. The close association of dark evergreen trees of medium size suggested the sclerophyllous Mediterranean forests described by European botanists (Figs. 84and85). Transition Salt Marsh to Pine Forest between Ocean Gate and Barnegat Pier Before reaching the fresh-water marsh, which was interspersed with wooded islands, so that the marsh ran between the islands in the form of Figure 82 Large white-oak, Quercus alba, at edge of salt marsh, Somers Point, April 6, 19 10, backed by a forest of Pinus rigida, Juniperus virginiana. Ilex opaca, fronted by Myrica carolinensis, Baccharis halimifolia, and Panicum virgatum. (Photograph by Henry Troth and J. W. Harshberger.) wet sloughs, a grove of Pinus rigida is entered with about 100 dominant trees. The undergrowth consisted of Acer rubrum, Diospyros vir- giniana, Nyssa sylvatica, Quercus falcata, and the shrubs Ilex glabra, Myrica carolinensis. Sassafras variifolium, and Vaccinium corymbosum, together with the herbs Gaultheria procumbens and Trientalis ameri- cana. The smaller islands were covered with medium-sized pine trees associated with Acer rubrum, Nyssa sylvatica, and an undergrowth of Andromeda mariana, Ilex glabra, Myrica carolinensis, Pyrus arbutifolia, 96 VEGETATION OF THE NEW JERSEY PINE-BARRENS Rhus copallina, R. radicans, Rosa Carolina, together with the ferns Aspidium thelypteris, Osmunda regahs, and the herb, Baptisia tinctoria. The sloughs between the islands were fringed with Panicum virgatum, Solidago sempervirens, which grew in drier ground, together with associations of the blue-flag. Iris versicolor, in pure growth, mingling at the water's edge with Castalia odorata, Hibiscus moscheutos, Os- munda regalis, Proserpinaca palustris, and grasses and sedges, such as Eleocharis tenuis and Scirpus Olneyi. The shrub which had advanced farthest into the marsh was Myrica carolinensis. Figure 83 Blending of salt marsh with Panicum virgatum at inner edge, Baccharis halimifolia, Myrica carolinensis, and forest with Pinus rigida dominant, fronted by Ilex opaca, Juniperus virginiana, Quercus alba, Q. stellata, Somers Point, April 6, 19 10. (Photo- graph by Henry Troth and J. W. Harshberger.) Pine-Barrens at Northern Limit Between Farmingdale and Allaire a small grove of Pinus rigida exists as a northern tongue of the main forest farther south. Here the pine trees are of all sizes, associated with Acer rubrum, Fagus grandifolia (as a low tree), Hicoria glabra, Castanea dentata, Prunus serotina, Q. alba, Q. rubra, Q. stellata, Q. velutina, and an undergrowth of such shrubs as Gaylussacia resinosa, Myrica carolinensis, Vaccinium corymbosum, V. stamineum, V. vacillans, with the herbs Chimaphila maculata and VEGETATION OF THE NEW JERSEY PINE-BARRENS 97 Figure 84 Salt marsh and pine forest, Somers Point, April 6, 19 10. Edge of the forest fronted with Myrica carolinensis, Juniperus virginiana (avi- vectent), ilex opaca, and a group of white- oal4 8 2.438 14 8.5 2.591 14 2-5 0.762 12 5 1.524 12 9 2-743 16 4 1.219 13 8.5 2.591 20 6 1.828 >4 6.5 1.981 18 6 1.828 1 1 8 2.438 13 7 2.133 19 6 1.828 9 3-5 1.066 1 1 2 0.609 9 AGE OF LARGER PINES GROWING WITHIN LIMITS OF L OWER PLAINS Height, Height, Age, Height, Height. Age, Height, Height, Age, Feet Meters Years Feet Meters Years Feet Meters Years 1 1 3352 30 I I 3-352 23 17 5.181 4> 10 3.048 13 '3 3.962 36 19 5.791 43 12 3657 16 13 3.962 40 21-5 6.553 41 11.5 3505 15 12 3.657 27 '7 5.181 33 12 3.657 24 12 3.657 28 10 3.048 22 20 6.096 40 11.5 3505 28 10.5 3.200 22 13 3.962 24 . 21 6.400 40 13 3.962 19 21 6.400 46 18 5.486 39 10 3.048 22 '7-5 5-334 42 ■7 5.181 38 * Pinchot, Gifford: The Plains. Annual Report, New Jersey State Geologist, li Forests: 125-130. 154 VEGETATION OF THE NEW JERSEY PINE-BARRENS AGE OF SPROUTS FROM THE UPPER, OR WEST, PLAINS Height, Height, Age, Height, Height, Age, Height, Height, Age, Feet Meters Years Feet Meters Years Feet M iTERS Years 4 1. 219 12 3 0.914 I I 7 2 ■33 '7 3 0.914 3' 2.5 0.762 7 3 0 914 7 7-5 2.286 22 4-5 1. 371 8 3-5 I 066 9 6 1.828 16 5 1.524 10 5 ' 524 10 5 1.524 18 5-5 1.676 16 4 I 219 1 1 8 2.438 13 4-5 1. 371 15 3-5 I 066 9 5-5 1.676 15 6 1.828 12 4-5 > 371 15 4 1.219 13 3 0.914 8 3-5 I 066 1 1 2-5 0.762 9 3-5 1.066 9 4.5 I 371 10 4 1. 219 8 6 1.828 13 3-5 ' 066 9 8 2.438 16 7-5 2.286 16 6 I 828 '5 5 1.524 >3 4 1. 219 7 4 • 219 10 6 1.828 15 5 1.524 12 5 ' 524 13 AGE OF LARGER PINES GROWING WITHIN THE UPPER PLAINS Height, Height, Age, Height, Height, Age, Height, Height, Age, Feet Meters Years Feet Meters Years Feet Meters Years 15 4.572 39 14 4.267 31 21 6.400 44 1 I 3352 21 9 2-743 22 13 3.962 33 12 3657 29 10 3.048 22 19 5791 46 14 4.267 30 16 4.876 30 '7 5. 181 39 I 1 3352 35 1 1 3352 36 '4 4.267 27 From the figures presented in the tables it is evident that the oldest and largest trees are under fifty years of age, and that the small growth averages about ten to fifteen years. Synecology of the Coremal Before we consider the cause of the present condition of the plains vegetation it is important to study that vegetation in detail from the standpoint of its synecology. We find associated with the dwarf bushy pines, loaded with cones, usually 6 to 9 decimeters (2 to 3 feet) tall (Fig. 129), Quercus marylandica, also stunted, but sometimes reach- ing above the pines by a growth which is 1.2 meters (4 feet) tall. The laurel, Kalmia latifolia, in full flower in early June, is sometimes 1.5 to 1.8 meters (5 to 6 feet) tall on the higher knolls of the Lower Plains. Dwarf post-oak, Quercus stellata, occurs more commonly on the Lower Plains than on the Upper, while the bear-oak, Quercus ilicifolia( = nana), is a low, prostrate, acorn-bearing bush, sometimes growing taller and more tree-like under more favorable conditions. The dwarf trees form VEGETATION OF THE NEW JERSEY PINE-BARRENS •55 Figure 130 View across the Lower Plain with character- istic growth of low oaks and pines. In the middle distance are the two wagons of the International Phytogeographic party which visited New Jersey on July 28-29, '9'3- (Photograph by George E. Nichols.) the conspicuous elements of the vegetation (Figs. 130 and 131), and between them grow, in different associations, Andromeda mariana, Comp- tonia asplenifoha, Dendrium (Leiophyllum) buxifolium, Gaylussacia frondosa, G. resinosa, Hudsonia ericoides, Ilex glabra, Kalmia angustifolia, Myrica carolinen- sis, Pteris aquilina, Smilax glauca [Little Plain], Tephrosia (Cracca) virginiana, Vaccinium pennsylvanicum, V. vacillans, and on the ground in bare places grow Epigaea repens, Gaultheria procumbens, and Pyxidanthera barbulata (in full flower April 8,1912). The bearberry, Arcto- staphylos uva-ursi, grows in ex- tensive radiate masses on flat, sandy stretches of the plains (Fig. 133). The herbaceous plants of the plains, as far as noted in the growing condition in the three seasons of the year, are: Andro- pogon scoparius, Chrysopsis mariana, Euphorbia polygonifolia and Melampyrum americanum. The reindeer-lichen, Cladonia rangiferina, grows on wide stretches of white sand. The most local and peculiar plant of the Lower and Upper Plains is the broom-crowberry, Corema Conradii, which grows in two general forms (Figs. 134 and 135). The first type of plant is one which grows in dense cushions, about 3 deci- meters (i foot) tall (Fig. 135), its color varying from light green through dark green to a rich brown color, distributed in clumps between the prostrate pine trees. That this is the typic form of the plant is indicated by the fact that on the island of Nantucket, where it grows in great abundance in several widely separated localities, it is always found in the cushion form, and of the dark-green color. Figure 131 Low growth of the pines and oaks in the Lower Plain, New Jersey, with members of the International Phytogeographic Excursion collecting the characteristic plants. July 29, 1913. (Photograph by George E. Nichols.) 156 VEGETATION OF THE NEW JERSEY PINE-BARRENS On April 8, 1912, Corema, for which this formation has been desig- nated the Coremal, was in flower, with its staminate and pistillate flowers on different plants. The flowers, which are dioecious, or polygamous, are in terminal clusters, each in the axil of a scaly bract, and with five or six scarious, imbricated bractlets, but no proper calyx. The stamens are three or four in number and exserted, with purple filaments and brownish-purple anthers. The style is slender, usually 3-cleft. The drupaceous fruit contains 3 (rarely 4-5) nutlets. The plants are low shrubs, with subverticillate, narrowly linear, heath-like leaves. In other places on the Lower Plains Corema assumes the second type of growth, which is a diffuse one, the separate stems being scattered widely. Such plants may be young plants that have arisen from scattered seeds. The local dis- tribution of this interesting plant early attracted the atten- tion of systematic botanists, and there is a copious literature which Stone* quotes in his book on the flora of the southern part of the State. Lately Brown has figured and described the plant in Bartonia,t but neither of these botanists describe the broom-crowberry from the syne- cologic point of view. The writer hopes a little later to emphasize, in an especially illustrated paper, some of the neglected points about the growth and distribution of this striking cushion-plant. In a basin-shaped depression in the West Plains Quercus marylandica forms a closed association, the trees growing 2.4 meters (8 feet) tall. The pitch-pine, Pinus rigida, is secondary to the oaks, and the under- growth consists of Comptonia asplenifolia, Gaylussacia resinosa, Kalmia latifolia and Quercus ilicifolia, while the low-growing species in these depressions are Arctostaphylos uva-ursi, Gaylussacia frondosa, Pyxi- danthera barbulata and Tephrosia virginiana. On an elevation nearly covered with a coarse gravel soil the trees become dwarfed, gnarled and twisted. Figure 132 Upper Plains (Little Plains) back of George Cranmer's farm. The tall tree is Quercus stellata. Note clumps of Corema Conradii with Cladonia rangiferina, April 9, 1912. * Stone, Witmer: Loc. cit.: 530-536. t Brown, Stewardson; Bartonia, 19 13 (No. 6): 1-7. VEGETATION OF THE NEW JERSEY PINE-BARRENS 157 A valley-like depression in the Lower Plains was filled with water and saturated sphagnum. The pines in these wet places were at least 4.5 - / 3 i- 1 : ;^^^^^^^^^^^?fa^^,. SBBlJ^sSS^Sety' '^ V '^-^^ ^^i^l ^s^^^w ^^^^^^^^^^ W^^^^^^^^^tM0^^^^^ I^^^^^^^^^^^^B ^jt^^^^^. ^ ..sp- , '^^S^&i jM^^^^^^^^^^^^^^^ w^ .^J^m^ '^^^^ ^^^^^^^^^^^^M Figure 133 Lower Plains with bearberry, Arctostaphylos uva-ursi, dwarf oaks, dwarf pitch- pines, and reindeer-lichen, Ciadonia rangiferina, April 9, 1912. to 6 meters (15 to 20 feet) tall ( red-maple, Acer rubrum, such shrubs as: Azalea viscosa, Clethra alnifolia, Ilex glabra, Kalmia angustifolia, Myrica carolinensis, Nyssa sylvatica, Vaccinium macrocarpon, the bracken-fern, Pteris aquilina, and the turkey's-beard, Xero- phyllum asphodeloides. The gray-poplar, Betula populifolia, is an interloper. The head of this small valley was charac- terized by eight tall pine trees forming a small grove com- pletely surrounded by low vegetation (Fig. 136). Here Figs. 136 and 137), associated with the Figure 134 Lower Plains showing prostrate pitch-pines and two clumps of Corema: the one to the left is green and staminate; the one to the right is chocolate-brown in color and pistil- late. Flowers present on April 9, 1912. 158 VEGETATION OF THE NEW JERSEY PINE-BARRENS edaphic conditions control the size and distribution of the pine trees. The Hne of demarcation between the plains and the pine-barren forest seems to be a sharp one, but nevertheless there are a few scattered tall pine trees at the outer margin of the plains which become reduced in height as the Coremal is entered. Here, in the forest of tall pine trees, the oaks are subordinated to the pines, which at the edge of the plains Figure 135 Corema Conradii as a cushion shrub along the road in the Lower Plain, N. J., July 29, 19 1 3. (Photograph by George E. Nichols.) are 6 to 9 meters (20 to 30 feet) tall, associated with stagger-bush, Andromeda (Pieris) mariana, sweet-fern, Comptonia asplenifolia, huckle- berries, Gaylussacia frondosa, G. resinosa, sheep-laurel, Kalmia angusti- folia, bear-oak, Quercus ilicifolia, and goat's-rue, Tephrosia (Cracca) virginiana. The vegetation of the plains joins sharply with that of a white-cedar swamp. A difference of 3 to 6 decimeters in soil level makes a marked difference in the vegetation along a stream's edge. Here the tension strip is characterized by Acer rubrum, Andromeda VEGETATION OF THE NEW JERSEY PINE-BARRENS I59 mariana, Chamaedaphne (Cassandra) calyculata, Clethra alnifolia and Myrica carolinensis. Measurement of Species of the Coremal In order to determine the actual size of some of the individual trees, shrubs and herbs characteristic of the Coremal the following measure- ments were made by a meter stick in the field. Trees Pinus rigida 40 centimeters to i meter tall. Quercus marylandica -f- i meter tall. Shrubs Arctostaphylos uva-ursi 10 centimeters tall. Corema Conradii 16 Dendrium buxifolium 50 " " Gaylussacia resinosa 40 " " ilex glabra 40 " " Kaimia latifoiia 30 " " Quercus ilicifolia 70 " " Herbs Andropogon scoparius 80 Chrysopsis mariana 40 " " Hudsonia ericoides 15 Theories Concerning the Coremal Several theories have been advanced to account for the dwarf character of the pines and oaks in the Coremal. These will be given first in detail and then the experiments will be described upon which a scientific con- clusion may be based. The first consistent attempt to explain the cause of the present condition of the plains is by GiflFord Pinchot. We quote as follows: "Some have attributed the form of the pine to a lack of mineral constituents in the soil necessary to the growth of trees. This theory, however, is disproved by chemical analyses of the soil made by the Geological Survey, which show no greater poverty than is common to the surrounding region. The theory that fire, combined with the effect of the very poor top-soil and a hard subsoil, is the efficient cause has been advanced and certainly fire has been a very large factor in bringing about the present conditions of these areas. If the plains have been in their present condition since the country was first settled, they were probabiy first burned over by the Indians, who were in the habit of camping in the neighborhood, as their shell-heaps show. It is not un- reasonable, therefore, to suppose that these high-rolling plains were originally stripped of their forest cover in this way. The pines, which probably returned by seed after the first fire, were burned over again and again, and their stumps sent up sprouts which became more and more i6o VEGETATION OF THE NEW JERSEY PINE-BARRENS feeble after successive fires. Many old stumps, as already pointed out, became exhausted and died after repeated sprouting. Their place was taken by seedlings and in this way the ground remained stocked with pine." Pinchot goes on to say: "This does not explain, however, the pros- trate form of the young seedlings and many older trees. This peculiar- ity is not confined to the plains alone, for pine seedlings growing on bare sand in exposed situations in the neighborhood of the plains show the same tendency. These seedlings exhibit a remarkable similarity to the forms assumed by trees near the timber line on high mountains. It is a fair inference that the very harsh and windy situations in which they grow has an effect analogous to that of a great elevation. Hence it is believed that exposure and poor soil are entirely sufficient to ex- plain why the young trees are prostrate. A large part of the pine, however, is coppice growth from old stumps which have lost their vigor to a large extent, and under the unfavorable surround- ings are incapable of producing anything but straggling sprouts. Furthermore, the trees, which grow very slowly on the poor soil of the plains, are killed by fire before they have time to reach a large size." George Cranmer, a farmer of Warren (Cedar) Grove, living on a farm at the edge of the Little Plain, showed me a portion of the plains back of his fields which he said had not been burned over for sixty years. The pine trees and the oak trees were taller on this tract and the growth of other species in every respect thicker, and yet, contrasted with the nearby pine forests, the trees of this protected area might with propriety be termed dwarf. This farmer ad- vanced the idea of soil exhaustion, viz., that on account of the poverty of the soil the trees had become poorer and poorer. The production of toxic excretions by the roots of the plants of the Coremal may be a factor of importance in inhibiting the growth of the pines and the oak trees found in this formation. Following out a clue which the partial sterilization of the soil by chemicals or by steam gave, it was discovered that the bacteria, which are useful in ammonia-making, Figure 136 Lower Plains showing group of tall pitch- pine trees at head of a wet hollow, April 9, 1912. VEGETATION OF THE NEW JERSEY PINE-BARRENS i6i increased four-fold after such treatment, suggesting the presence in the soil of some agent which held them in check. After much painstaking study it was discovered that the soil contained a living protozoan (Pleurotricha), which preyed upon the useful organisms, and that the heat and chemicals either destroyed these larger unicellular animals or inhibited their activity. It can be said, therefore, when such cases are considered, that the fertility of the soil may be in part a biologic problem.* That the productivity of some soils is due to biologic rather than to physic and chemic characteristics is illustrated by the attempts made to reforest Denmark. The peninsula of Jutland was covered originally by forests, but these were destroyed, until, by the year 1500, the country had been transformed into a barren heath and sand dunes. At various times attempts were made to reforest these heaths, but the results were disappoint- ing until Colonel E. Delgar solved the problem. Spruce trees (Picea alba, P. excelsa), if planted alone, did not thrive, but became sickly. The cause of this irregularity in the growth of spruce was thought to be local conditions of the soil, but scientific investigation of such soils did not reveal any differ- ence in the physic or chemic com- position of the soil, it was found, however, that the mountain pine (Pinus montana) acted as a nurse to spruce trees planted in its vicinity. In the same soil where spruce if planted alone would remain backward, it would, if planted close to a mountain pine, grow up vigorously. After some years of trial it was found that the pine would hamper the growth of the spruce, and so it was cut down at an early age. It was discovered then that even if the mountain pine was cut down at an early age, it imparted to the adjacent spruce trees the ability to grow. The phenom- enon is not understood, but it is supposed that the roots of the mountain pine are inhabited by some mycorhiza which produces the nitrogen necessary for the growth of trees, and that this organism is transferred to roots of the surrounding spruce trees. Once this infection has taken Figure 137 Lower Plains, showing a sedgy slough sur- rounded by pitch-pine trees (scattered), April 9, 19 12. Harshberger, J. W.: The Soil: A Living Thing. Science, new ser., xxxiii : 741- 744- I 62 VEGETATION OF THE NEW JERSEY PINE-BARRENS place, the presence of the mountain pine is no longer necessary and it is usually cut down. Clearly this is a biologic relationship. The thought arises naturally with reference to the soil of the plains, whether the tall pines can be grown there, if a nurse crop is discovered similar to the mountain pine above described. It seems plausible that the poverty of the soil in the Upper and Lower Plains is due to the de- velopment of toxic substances which might be overcome by planting trees that are indifferent to such soil toxins, as has been accomplished so admirably in Denmark. Future investigation of these soils will alone determine these facts. Figure 138 Culture of corn in soil from Upper (Little) Plains to test character of top- and subsoils. Left, top-soil untreated; middle, subsoil untreated. Right, garden soil as control. (Photo- graph taken May 20, 1912.) '%^\ ^ " v ^ Figure 139 Culture of peas in soil from Upper (Little) Plains to test character of top- and subsoils. Left, top-soil untreated. Right, subsoil untreated. (Photograph taken May 20, 1912.) Experimental Treatment of Coremal Soils Realizing that the problems connected with the vegetation of the plains of New Jersey and with their soils lent themselves to experimental in- vestigation, the writer began a series of experiments to test the character of the soil: For this purpose a box of soil from the Little Plains was pro- cured. The samples included subsoil and top-soil. These were placed in pans and flower-pots with and without previous sterilization by heat. In the sterilized and unsterilized top-soil and subsoil corn, peas, sun- VEGETATION OF THE NEW JERSEY PINE-BARRENS 163 Figure 140 Culture of corn in series of soils from Upper (Little) Plains to test character of soils. Left, top-soil untreated. Second left, top-soil heated; subsoil untreated; subsoil heated. Right, garden soil untreated; garden soil heated. (Photograph May 20, 1912.) flower, wheat and okra seeds were planted on April 13, 1912, and placed in a cold-frame out-of-doors under as natural conditions as possible (Figs. 138, 139, 140, 141, 142). Nine days after the seeds were planted, on April 22, the pots and pans in the cold-frame were examined. In all of the soils, treated and untreated, the seeds had sprouted and their plumules were seen projecting above the surface in the case of the peas, sunflower and wheat. Two days afterward, on April 24, the peas, sunflower and wheat pushed their plumules well above the soil. Prog- ress had been made in the growth of the seedlings of peas, sunflower and wheat, but at this early date there was seen an inhibition of growth in the subsoil, treated and untreated. Subsequent to April 29, when the corn-tips began to show above ground, the young plants in the pots and pans containing subsoil, treated and untreated, showed a constant lagging behind, due perhaps to the stiff, com- pact nature of the soil in which the corn was planted. The details of the experi- ments, as recorded in the notebooks, are too volumin- ous for complete description here — only the general re- sults can be presented. The experiments ran thirty-eight days, when some of the pot- grown plants were lifted and placed between felt driers for preservation (Figs. 138, 139, 140,141,142). The contrast in size and growth was best seen in the shallow pan cultures. Contrasting the three pans of corn with top-soil untreated, the plants were 12.7 centi- meters (5 inches) tall, leaves fully expanded, but slightly yellowed; Figure 141 Culture of peas in series of soils from Upper (Little) Plains to test character of soils. Left, top- soil untreated. Second left, top-soil heated; subsoil untreated; subsoil heated; garden soil untreated. Right, garden soil heated. (Photograph taken iMay 20, 1912.) 164 VEGETATION OF THE NEW JERSEY PINE-BARRENS subsoil untreated, plants about 12.7 centimeters (5 inches) tall, yellowish in color, leaves suberect and inrolled; garden soil, plants 12.7 to 17.7 centimeters (6 to 7 inches) tall, leaves broadly expanded, dark green. Contrasting the two pans of peas with top-soil untreated plants 20.3 to 25.4 centimeters (8 to 10 inches) tall, the peas were more withered and semi-prostrate, the soil less retentive of moisture; subsoil untreated, plants more erect, less wilted, darker green; soil stiffer and more reten- tive of water. It will be seen from the photographs taken of the soil cultures on April 10, 191 2, that the plants grown in the stiff, heavy, impervious subsoil are of small size, more starved in appear- ance and show every evi- dence of sterile conditions of soil environment. The influ- ence of the sterile subsoil is illustrated best in the pot cultures of corn, wheat and sunflower and least pro- nounced in that of the peas. The height of the plants is dwarfed, the secondary roots are shorter and the plants have an underfed appearance. The sandy top-soil of the plains, although containing some humus and relatively porous compared with the impervious, hard-baked subsoil, was less decidedly rich than the garden soil used as a control. The plants of corn, wheat and sun- flower were smaller than those Figure 142 Cultures of wheat and sunflowers in series of soils from Upper (Little) Plains to test character of soils. Top set, left top-soil untreated; top-soil heated; subsoil untreated; subsoil heated; gar- den soil untreated. Right, garden soil heated. Lower set, sunflower plants, series of soils same as in top set, and arranged in same order. grown in ordinary garden soil, and the better growth was made in the unheated or unsterilized top-soil, as contrasted with the heated or ster- ilized soil (Figs. 143, 144, 145, 146, 147)- From these experiments and the series of control experiments with soils from the pine forest just over the west edge of the plains and in which normal growth was secured in top- and subsoils (see figures), we are probably safe in assuming that the inhibition of the growth of the va- rious garden plants, including the wild plants of the plains, is due to the impervious, stiff and easily baked subsoil through which the roots can VEGETATION OF THE NEW JERSEY PINE-BARRENS 165 hardly penetrate, and which supplies very little plant food, but that the content of the soil in living organisms may have some influence is indi- cated by the difference of growth between the plants raised experimen- tally in heated and unheated soils (Figs. 148 and 149). That with the application of proper fertilizer, the top, or light, sandy soils of the plains could be made to produce fair crops of corn and wheat is indicated on Cranmer's farm, adjoining the plains, where corn, pumpkins, tomatoes, beets, cabbages and sweet potatoes are grown, but owing to the nature of the subsoil these attempts at cultivation will meet with very little commercial success as long as the stiff bottom soil remains as it is. The roots of the pines and the oaks, as indicated by digging them up, cannot penetrate Figure 143 Culture of corn continued to July 4, 1912, in open pans with subsoil untreated. Figure 144 Culture of corn continued to July 4, 1912, in open pans, when photograph was taken; top-soil untreated. any distance into this hardpan. They decay after making futile at- tempts to penetrate the subsoil saturated with cold water and cemented together into a compact mass. The plants of the plains, with short, superficially formed roots, get along fairly well in the pervious sandy top-soil. Some facts may be learned about the vegetation of the New Jersey Coremal by contrasting it with the Coremal on the Island of Nantucket. The physiognomy of the vegetation is nearly alike on the Upper and the Lower Plains of New Jersey and on the Island of Nantucket, but the Nantucket Coremal (Fig. 150) lacks the pitch-pine. In both districts 1 66 VEGETATION OF THE NEW JERSEY PINE-BARRENS there is a low vegetation of shrubby oaks, Ouercus ilicifoha ( = nana) and Q. prinoides; the bearberry, Arctostaphylos uva-ursi, is more abundant on the Island of Nantucket than on the plains of New Jersey, while the broom-crowberry, Corema Conradii, is found in both districts. A com- parison of the Nantucket vegetation gives a clue to the origin of the plains (Coremal) and pine-barren vegetation of New Jersey. Heath- land is the result of the factors which are summed up under the general term oceanic climate, and Nantucket, isolated as it is far out at sea, has an oceanic climate. The strong winds that blow and the pervious glacial soils prevent the deciduous trees of larger size from spread- ing out of the valleys, where they are protected, and even the pitch-pine, recently intro- duced into Nantucket, follows the valleys and protected slopes of the hills (Fig. 151). In all probability when the pine-barren region of New Jer- sey was an island an oceanic climate prevailed. The heath- land, of which the Coremal is a part, was left as a relict as the plains of New Jersey. The pine-forest in those early times probably filled the valleys and later spread over the hills until all of the region was covered with pine forest except the Up- per and Lower Plains, where edaphic conditions prevented the development of tall pine trees. The heathland was converted into a pine-heath. Graebner* has detailed a similar conversion of heath into pine-forest by the invasion of pines, and in such pine forests the undergrowth consists of characteristic heath plants, hence Kiefernheide. The reason why the pine trees have been unable to grow to tall size in the New Jersey plains is because of the hard layer of soil immediately below the upper sandy layer. This layer corresponds to the caleche of Mexico, the plow-sole of agriculturists and the Ortstein of the Germans. *Graebner, P.: Die Heide Norddeutschlands. Vol. V. Die Vegetation der Erde: 239. Figure 145 Culture of corn continued to July 4, 1912, in open pans, with garden soil untreated. VEGETATION OF THE NEW JERSEY PINE-BARRENS 167 Klebahn* figures an Ortstein-Kiefer where the tap-root striking the hardpan is bent over, being unable to penetrate this soil layer. Graebner narrates how such pine trees grow for a time, but finally, after reach- ing a certain age, begin to go back until they succumb, and he de- scribes how certain pine trees more fortunately situated by natural planting over holes through the Ortstein (Ortsteintopfe) are able to send their tap-roots into the deeper soil layers. t Under such conditions tall thrifty pine trees will be scattered here and there, while the majority of the trees that become established in the region are dwarf and languish- ing. Similar conditions are found in the plains of New Jersey, where the 1 *«,•!<' 'U^M--' 'i' ' ■ / Figure 146 Culture of sunflowers continued to July 4, 1912, wfien photograph was taken. Left, top- soil untreated; top-soil heated; subsoil untreated; subsoil heated. Right, garden soil un- treated. low, dwarf pine trees live for a number of years and finally succumb, to be replaced by other trees that pass through a similar existence. Thus hardpan and fire are the two most important factors which have per- petuated the heath vegetation of the plains, while the surrounding region, with more pervious soil, although similarly fire swept in later years, has been preserved as a pine forest or pine heath (Fig. 151). Remove the pines and you would have the conditions as they exist on the Nan- tucket heathland and the vegetation would have a similar physiognomy, with such low oaks as Quercus ilicifolia and Q. prinoides forming the * Klebahn, H.: Grundzuge der allgemeinem Phytopathologie: 14. t Graebner, P.: bottom of page 125. 1 68 VEGETATION OF THE NEW JERSEY PINE-BARRENS main ground-cover (Fig. 150). We are able, therefore, by this com- parative study to connect three interesting plant formations with those of Europe. America Nantucket Heathland (with low oaks and bearberry, Arctostaphylos). New Jersey Coremal (heathland with low pines, etc., in Lower and Upper Plains). New Jersey Pine-barrens. Europe Oak Heath {Eichenhcide) (with Arctostaphy- los Heath, /. e., Heide mit Vorherrschen von Arctostaphylos). Low Pine Heathland (Kiefernheide). Pine Heathland ( Kiefernheide). Summing up our study of the Coremal, or the plains vegetation, we may state that the dwarfed character of the pines, oaks, and other plants Figure 147 Culture of corn and wheat continued to July 4, 19 12, when photograph was taken. Left, corn in top-soil untreated; garden soil untreated. Wheat, left, top-soil untreated; subsoil untreated; subsoil heated; garden soil untreated. of the plains is due primarily to the stiff, impervious subsoil, and the light, easily dried-out, sandy surface soil which may freeze to the depth of a meter in winter and dry out in midsummer until the subsoil is hard and stiff and the surface soil almost devoid of moisture. The ele- vated character of the country facilitates rapid drainage by run-off and seepage, and this influence of elevation, coupled with the strong winds that blow, has a strong dwarfing effect on the plants exposed to such conditions. These factors are sufficient to account for the origin of the Coremal as a distinct formation. Fire has had an influence on the native vegetation, but its effect has been secondary to the soil factors, for forest fires have burned over as severely and as repeatedly the nearby forest areas without so appreciably stunting the growth of the pine trees. VEGETATION OF THE NEW JERSEY PINE-BARRENS 169 Figure 148 Culture of corn in top-soil (left) and sub- soil (right) from pine forest immediately west of the Upper (Little) Plains, started as a control of studies in the character of Upper Plains soils. Undoubtedly it is true that the pines naturally of dwarf size in the Core- mal, after having been burned again and again, have sent up from the charred stumps more and more feeble sprouts until many old stumps have become exhausted and have died from loss of vigor. The areas of the plains, then, are those which have edaphic- ally an unusually impervious subsoil. In the deposit of this subsoil we can conceive of its being laid down as a sheet, or covering, of relatively im- pervious material of circumscribed area, surrounded by soils in which the subsoil is more favorable to the deeply penetrating roots of the pitch- pines and other pine-barren trees, as the writer has proved by ample con- trol experiments. After deposit these impervious areas of subsoil were cov- ered by more pervious deposits of light sands and gravels. The plains are located as they are geographic- ally because it was in parts of Ocean and Burlington counties that the materials out of which the stiff sub- soils were later formed were depos- ited. The factors responsible for the development of the succession which we have termed the Coremal, or plains formation, are soil, elevation, wind, and fire. Fire has played a secon- dary part in the development of the physiognomy of the vegetation. One of the strongest proofs that the plains have existed for many successional periods as such is the presence and persistenceof Corema Conradii, which has remained as a characteristic plant of the Coremal ever since the time when, as in Nantucket, the pine- barren region of New Jersey had an Figure 149 Culture of peas in top-soil (left) and sub- soil (right) from pine forest immediately west of the Upper (Little) Plains, started as a control of studies in the character of the soils of the Upper Plains. 1 70 VEGETATION OF THE NEW JERSEY PINE-BARRENS oceanic climate when isolated as an island from the mainland by Pen- sauken Sound. Vegetation of the Hempstead Plains, Long Island* In order to have a slightly different viewpoint in our study of the plains vegetation of New Jersey a short account of the vegetation of the Hemp- stead Plains in western Long Island will be given. It is the belief of the writer that the Hempstead Plains represent one element in the series of Figure 150 Nantucket Heath back of Sankaty Head, covered with a dense low growth of the dwarf chestnut-oak, Quercus prinoides, August 25, 1913. heathlands which include the plains of New Jersey and the heathland of Nantucket, for the floristic elements in all three of these formations are much the same. In all probability they originated in the same epoch of geologic time. The country covered by the Hempstead plains vegetation is fiat and resembles the western prairie plains both in general physiognomy and in the association of plant species. The vegetation is of the grassland * Harper, Roland M.: The Hempstead Plains. Bull. Amer. Geogr. Soc, xliii: 351- 360. VEGETATION OF THE NEW JERSEY PINE-BARRENS 171 type rather than the heathland type. The plain is practically treeless, although dotted over with low trees, it is bisected by the valley of Meadow Brook, which flows south into the Atlantic Ocean. The scat- tered dwarf trees of the western half of the plains are the white birch, Betula populifolia, and the aspen poplar, Populus tremuloides, both of which are distributed by wind-carried fruits. The eastern half possesses, in addition to these two trees, occasional dwarf pitch-pine trees, Pinus rigida, and dwarf-oaks, such as Quercus ilicifolia, Q. prinoides, Q. stellata, all of which are scattered as individuals or exist in rounded clumps. The shrubs comprise, according to my notes, the stagger- bush, Andromeda (Pieris) mariana, sweet-fern, Comptonia asplenifolia, huckleberry, Gaylussacia resinosa, Rhus copallina, and a low willow, Salix humilis. The most abun- dant and important prairie plant is the beard-grass, Andropogon scoparius, which covers many parts of the Hempstead Plain with an exclusive growth of its characteristic tufts or bunches, so that the general physiognomy is that of a western bunch-grass prairie. The most pronounced bunch-grass prairie effect is on a gravelly hillside along East Meadow Brook. Here Andro- pogon scoparius is undisputed possessor of the gravelly slopes, and with it a few herbs are found: Asclepias verticillata, Baptisia tinctoria, patches of reindeer- lichen, Cladonia rangiferina, and the rush, Juncus Greenei, and in the spring masses of Viola pedata. Several areas in western Nantucket depart from the prevailing heather vegetation and assume the physiog- nomy of the prairie. The beard-grass, Andropogon scoparius, is the pre- vailing grass in such grassland. It will be noted, from the above description, that the physiognomy of the vegetation and the floristic character of the associated species are different from those described for the Lower and Upper Plains vegeta- tion (Coremal) of New Jersey in the almost entire absence of low trees and in the presence of the prevailing grass, Andropogon scoparius, which gives characteristic tone and color to the Hempstead Plains of Long Island. Fk.l'ki-; 131 Inland sand-dunes, Nantucket, covered by marram-grass, Ammophila arenaria, and in- vaded by the pitch-pine, Pinus rigida, with Hummoci< Pond at the left. August 24, 1913. CHAPTER XII CULTIVATED PLANTS OF THE PINE-BARREN REGION A consideration of the vegetation of the New Jersey pine-barrens would not be complete if a brief consideration of the cultivated plants which have been grown successfully were omitted. The character of the successful garden and field plants is an index of the soil conditions and of the climate under which such plants are grown. A foreign phytogeog- rapher can appreciate better the soil and the climatic conditions by referring to a list of the common cultivated plants of a region than in any other way. To indicate that certain cultivated species thrive or do not thrive in a district is a way of stating that the soil and climate in general are not suitable for such crops. Three sources of information have been used in the preparation of the following list. Through the courtesy of William F. Bassett, nurseryman and botanist of Hammonton, New Jersey, and Robert D. Maltby, Dean of the Baron de Hirsch Agri- cultural School, Woodbine, New Jersey, I am able to supplement my list of the cultivated plants of the pine-barrens by some additional ones. Forest Trees White pine (Pinus strobus). Austrian pine (Pinus sylvestris). Black walnut (Juglans nigra). Butternut (Juglans cinerea). English walnut (Juglans regia). Pecan (Hicoria pecan). Locust (Robinia pseudacacia). Silver maple (Acer saccharinum). The forest trees in the above list are cultivated either for ornament or shade, and are not spontaneous, as the trees mentioned under the head of weeds. Fruit Trees Apple (Pyrus malus). Cherry (Prunus avium, P. cerasus). Peach (Prunus persica). Pear (Pyrus communis, P. sinensis). 1 um I (-pj-uj^yg cerasifera, P. domestica, etc.). Quince (Cydonia vulgaris). 172 VEGETATION OF THE NEW JERSEY PINE-BARRENS 173 Bush Fruits and Vines Blackberry (Rubus nigrobaccus). Currant (Ribes rubrum). Fig (Ficus carica). Gooseberry (Ribes grossularia). Grape (Vitis — several species and hybrids). Raspberry (Rubus idaeus, R.occidentalis (black)). Small Fruits Cranberry (Vaccinium macrocarpon). Strawberry (Fragaria chiloensis). Root Crops Onion (Allium cepa). Potato (Solanum tuberosum). Radish (Raphanus sativus). Sweet Potato (Ipomoea batatas). Turnip (Brassica rapa). Leaf Crops Cabbage (Brassica oleracea) Celery (Apium graveolens). Lettuce (Lactuca sativa). Fodder and Cereal Crops Alfalfa (Medicago sativa). Buckwheat (Fagopyrum esculentum). Corn (Zea mays). Cowpea (Vigna catjang). Millet (Setaria italica). Oat (Secale cereale). Sorghum (Sorghum vulgare). Wheat (Triticum vulgare). Flower Crops Aster (Callistephus, several species). Cosmos (Cosmos bipinnatus, C. diversifolius, C. sulphureus). Dahlia (Dahlia variabilis, etc.). Gladiolus (Gladiolus, several species and hybrids). Paeony (Paeonia, several species). Sweet Pea (Lathyrus odoratus). 174 vegetation of the new jersey pine-barrens Garden Crops D / Lima (Phaseolus lunatus). t^ean j ^^^ (Phaseolus vulgaris). Canteloupe (Cucumis melo var. cantelupensis). Cucumber (Cucumis sativa). Okra (Hibiscus esculentus). Pea (Pisum sativum). Peanut (Arachis hypogaea). Pepper (Capsicum annuum). Pumpkin (Cucurbita pepo). Soy (Soja hispida). Tomato (Lycopersicum esculentum). CHAPTER XIII GENERAL OBSERVATIONS ON PINE-BARREN VEGETATION Within recent years a number of European botanists have attempted to take a census of the vegetation of phytogeographic regions by counting the number of plants found in a number of limited areas, diversely situ- ated as regards their habitats, and from the figures thus obtained to estimate the plant population of that region. Approximate values are obtained, which are useful in the comparative study of the vegetation of other regions. Clements,* in this country, advocates the use of a quadrat. A quadrat is merely a square of varying size, marked oflf in a formation for the purpose of obtaining accurate information as to the number and grouping of the plants present. Thus, by the use of the quadrat, it was ascertained that 259 hectares (one square mile) of alpine meadow contained approximately 1,500,000,000 plants. A more ex- tended use of the quadrat has been to determine the relative rank of the species in a formation, the seasonal aspects of the vegetation, and by means of permanent and denuded quadrats to determine the seasonal changes in a formation and the general mode of invasion of species into a quadrat deprived of all growing plants. Jaccard has applied somewhat the same method to a study of the flora of the Alps and the Jura. The papers received by me from this botanist in the order of their publication are: Etude Comparative de la Distribution Florale dans une Portion des Alpes et du J ura. Bulletin de la Societe Vaudoise des Sciences Naturelle, 4e S., vol. XXXVII, No. 142 (1901). Gesetze der Pflanzenvertheilung in der alpinen Region auf Grund statistisch-floristischer Untersuchungen, Flora, 1902, 111. Heft, 90 Bd. Nouvelles Recherches sur la Distribution Florale. Bulletin de la So- ciete Vaudoise des Sciences Naturelle, ^ P^r cent. The generic coefficient is emphasized by Jaccard as of importance in a comparative study of vegetation. This is obtained by contrasting the number of genera and the number of species of a flora. For example: the entire alpine region of the southern Jura has 54 genera to a 100 species; the whole Swiss flora has 26 genera to a 100 species. The generic coefficient of the alpine Jura is 54 per cent., and of the Swiss flora, 26 per cent. Jaccard has studied also the individual frequence (Frequence individuelle), the floral density (Dichte eines Bestandes = densite florale), and other statistic questions concerning the alpine flora. His third paper is embellished by several important graphic tables. Oliver and Tansley,* English botanists, have applied their methods of surveying vegetation on a large scale by a study of a salt marsh occu- pying the floor of an estuary called Bouche d'Erquay, on the north coast of Brittany. The method employed by them may be termed the "Method of Squares" and the "Gridiron Method" respectively. The former is suitable for the purpose of constructing a general map of an area comprising a considerable number of acres, on a scale of yIo to 5^0 or thereabouts, the latter for very detailed work restricted to small parcels of ground and giving results correct to six inches or even less. The method of squares is to peg out a square on the ground, which can be quickly and accurately mapped upon sectional paper. The grid- iron method based on the first is suitable where the physic features with which the different plant formations are correlated exhibit definite variations within quite short distances. Pine-Barren Vegetation by Quadrats The method adopted in the statistic study of the New Jersey pine-barren vegetation is a slight modification of the method of squares, or quadrats. A selected area of pine-barren vegetation was inclosed within a square with ten-meter sides with stakes driven in at the corners, and at inter- vals of a meter along the sides. Six lengths of white binding tape were used, each 10 meters long. The meter intervals were indicated by a piece of red string tied around the tape. The large square was first located by stretching the four lengths of tape, and the smaller meter * Oliver, F. W., and Tansley, A. G.: Methods of Surveying Vegetation on a Large Scale. The New Phytoiogist, iii: 228-237, November and December, 1904, with plate .XI and Figs. 77-80. VEGETATION OF THE NEW JERSEY PINE-BARRENS •77 squares were located by moving the other two pieces of tape as the survey proceeded, placing them parallel between two adjacent meter stakes on both sides of the outer and larger square. Thus each pine-barren area surveyed included the detailed study and mapping of lOO quadrats, each a square meter. As the method is a laborious one, requiring nearly a whole day's work on the hands and knees in a country infested by mosquitoes, only three of the large squares were finished. uM M MMe. ^ M MMm M ^^ ^ MM M M ^ ^ «\.\« « ' X MM MMM » <3 X' X X MM IJIw X m" "^ K " M M ° M "m m M W.I. fiXi « X xx" "m- X X q' " M.MMM ''M^^M M " X J^ M M M x^M'^ MM-M e ■^ MM * a,MMm M M M M M » X "\ ' '« p M m" fMHt-J ^ J & Q «^ ^M ^eM 4 M 1.^ < ;• M M Mm ^ ^ 0^ K-:* ^M M. •♦ ^♦Jt MQ M Q Q 0 q q ■^ ff MM M M M '.*. tfM M M ^ ■» nn^ T M M ' 6 t 4 0 ¥ yZ m q n, ^ q q, P S SS q q k|3 "m m" - A ••■. ft f * " t •p " 8 em mm ». i:^^ o Q Q S"- _M_ tit + Figure 152 Quadrat i, north side of d, Amelanchier intermedia. p, a, Ampelopsis quinquefolia. &, A, Asclepias obtusifolia. q, e, Euphorbia ipecacuanhae. Q ■f. Gaylussacia resinosa. $, M, Melampyrum americanum. '9', X, Moss. S, m, Myrica carolinensis. s, if, Pinus rigida. v. Como Lake. Prunus serotina. Quercus alba. Quercus ilicifolia. Quercus marylandica. Quercus prinus. Quercus stellata. Smilax glauca. Solidago odora. Vaccinium pennsylvanicum. The first survey was made in the pine woods on the north side of Como Lake on August 23, 1910, in the coastal pine-barrens (Fig. 152). The second was made in the pine forest 1.6 kilometers (i mile) east of Lakehurst, in the heart of the pine-barrens (Fig. 153), and the third quadrat was located 0.8 kilometer (}4 mile) east of Sumner, at the west- ern edge of the pine-barrens (Fig. 1 54). These three localities represent a wide diversity of soil, climatic, and vegetational conditions. The data 178 VEGETATION OF THE NEW JERSEY PINE-BARRENS for an estimation of the plant population of the pine-barren is based on the total area of the region comprising 1,200,000 acres* out of 4,988,800 acres (7,795 square miles), the total area of the State of New Jersey. The pine-barren region is, therefore, approximately 25 per cent, the total area of the State. As one acre is equivalent to 4,046.87 square meters, the total area of the pine-barren region is 4,856,144,000 square meters. ySygs KVvQc Va*„ „ »« e \raa o o oo qVv'S ^ v V « GG w£ V ^, ^,■«^»g Cy B|cg8«VVj» V«c «)f^OVg^ W V V e g y^c c °VVV r'VVKQVVV "v* gccG c ^KkVqSV ^..IJgt f V V q Ve vVj V Vgq g .3^:: V K KCc V„ q to V « « svc°c V t f q V V *gVgk' V q -g^R ftjV ^'^rV^VgC V«l°»et ^v'JiKf^^ V Q " , 0 c y '^gq V * " ^v'' g * •" g t r g f t V ^v q g e w VG V Vgv gv rn^v/sKK V vv* 1 q^'eVgv'g V cg c ,v v g ^g ' s g v »rQ ►d' v*«X«v» » >; 1 c Vg ^fV.f'i: l^'i^^^y A ^x ft V fVQ\ V^l V C V (, 8 ^ >S/IVgV <■ «. ff f- "gS* *v> »t* VV>t>f "gf = ^1. V 4, vgv«> v;/!j„u^ V V »*«*w w> 8 tv** v'» g *>VvVIg V VV;„VVVVIq ^y/Vv t^r'^Xr ^S" q *,« gv^ ^*L gV*v* 8 a, ly V V VW" V VyJ' 1 V aI^* '' f V V 4* G «J o q , 8 „V ' V / ^ V&8^ tv/tVS 'M/VVA*Oif<'tV « 8 ♦•V \'T^ ' 1"56V'-VSV Figure 153 Quadrat 2, one mile east of Lakehurst. ^ A, Andromeda mariana. £, Quercus ilicifolia. c, Comptonia asplenifolia. Q, Quercus marylandica. g. Gaultheria procumbens. q, Quercus prinoides. Gaylussacia frondosa. X, Quercus tinctoria. ■f , Gaylussacia resinosa. I, Ilex glabra. M, Melampyrum americanum. it, Pinus rigida. P, Pteris aquilina. Sassafras variifolium. Smilax glauca. Tephrosia virginiana. V, Vaccinium vacillans. In the first (Como) ten square meter quadrat there were 1 5 pitch-pine trees. In the second ten square meter quadrat at Lakehurst there were 16 pitch-pine trees. In the third ten square meter quadrat at Sumner there were 15 pitch-pine trees. Taking 15 trees as the average for the three stations, and multiplying by ^^-^^^^^^^ = 485,614,400, we get as * Vermeule, C. C. gist, 1899: 1,6. The Forests of New Jersey. Annual Report of the State Geolo- VEGETATION OF THE NEW JERSEY PINE-BARRENS 179 the approximate total number of pine treesin the pine-barren region the astounding figure 7,284,216,000 trees of the pitch-pine, Pinus rigida. Of course these figures are only approximate, because the cleared areas have not been deducted, the areas denuded of their timber and the open coun- try of swamps and savannas have not been subtracted, but the figures are interesting as an approximation to a total census of the pine trees in the '*\^ t e k k P •> ' "vP «*v i^Pj vV "*'«!< KgC g"cPtC kt.t^'-k^ kk k "k "^ "' " t. "; ^v'' g'yVV, ►'v'/l ^y 'si" "Vp*P - k 1 **2f V v^ ■ "^ w t* v"! ■'v t \ ^ " JIfPV *"'PP^ f, ♦ ">- vVkki.' iff k ^; ^ ',p>^t' *Sp V ^Vve- >- •*"•' Vvt' ,;V ';'. «.;, *k"/ ^'v'''^ -e-^a i:'V8 V;vv>J! ku^-''ki\,'' V /ftp ^ ,'- " •* V ogHfck*!, ^V kky '^pt 1 V Vy V„ '■k KK I; ^ .. ^ u . " ., k y i» v* Pi o y y. yf-^ ^Vy ^ ''*'• KK P Figure Quadrat 3, one-half mi Clethra ainifolia. Comptonia asplenifolia. Epigaea repens. Gaultheria procumbens. Gaylussacia resinosa. Hicoria glabra. Kalmia angustifolia. Kalmia latifolia. Pinus rigida. Pteris aquilina. 154 le east of Sumner. Pyxidanthera barbulata. Quercus ilicifolia. Quercus prinoides. Quercus stellata. Sassafras variifolium. Smilax glauca. Toadstools. Vaccinium pennsylvanicum. Vaccinium vacillans. region. Similarly, using the three ten square meter quadrats as a basis, we can compute the total number of plants that approximately occur in the region. Quadrat i (Fig. 152), located in the coastal pine forest on the north shore of Como Lake, had 680 plants, which would make on the average 6.8 plants to each secondary quadrat, one square meter in dimensions. Quadrat 2 (Fig. 153), located in the heart of the pine- barrens one mile east of Lakehurst, had 2,071 plants, or 20.071 plants to each secondary quadrat one square meter. Quadrat 3 (Fig. 154), meas- l80 VEGETATION OF THE NEW JERSEY PINE-BARRENS ured in the pine forest at Sumner at the western edge of the pine-barren region, had 2,544 plants, or 25.44 plants to each secondary quadrat. The numbers 6.8, 20.071, 25.44 represent the floral density of each of the three quadrats. It is interesting to note that the smallest number of plants per secondary quadrat is found in the more sterile pervious soils of the coastal pine-barrens. As a corollary of this fact, we find that in arid districts the native shrubs and trees stand far apart, so that each individual tree has a large mass of soil from which to draw water and food. The thin character of the vegetation in the coastal pine-barrens is a direct expression of this law. The true pine-barren forest at Lakehurst has more favorable soil conditions, as reference to the dis- cussion of soils will indicate, while the vegetation of the western edge of the pine-barrens in close proximity to the deciduous forest has still a larger number of plants per quadrat. If we draw an average of the total number of plants in each of the three quadrats ten square meters in size, we fmd it to be 1,765 plants. Multiplying 1,765 by 485,614,400, the area of the pine-barrens expressed in ID square meters, we get as the total number of pine-barren plants, including mosses, fleshy fungi, herbs, shrubs, and trees, 857,109,416,000 plants. Deducting the number of pitch-pine trees, 7,284,216,000, from the above total we get 849,825,200,000, which represents the approximate total number of all other kinds of tree, shrub, herb, and lower plant found in the region. In round numbers, the pine trees represent 0.81 per cent, of the total number of plants in the whole region, and yet, con- sidering the size and bulk of the pine trees, they form the dominant and most conspicuous element of the vegetation. No attempt has been made to calculate the individual frequency, but the following statement will prove of value in drawing a comparison among the three quadrats and in estimating the actual occurrence of the several plants in the pine-barren region as a whole. Plants Common to the Three Quadrats Pinus rigida Quercus stellata Smilax glauca Vaccinium pennsylvanicum J Plants Common to Two Quadrats (Indicated by Numbers) Comptonia asplenifolia 2,3 Gaultheria procumbens 2, 3 Gaylussacia resinosa 2, 3 Melampyrum americanum 1,2 Pteris aquilina 2, 3 , ^^ Quercus ilicifolia 1.3 " marylandica 1,2 " prinoides 2, 3 Sassafras variifolium 2,3 Vaccinium vacilians i> 2 VEGETATION OF THE NEW JERSEY PINE-BARRENS Plants Found in One Quadrat Only (Indicated) Amelanchier intermedia i Ampelopsis quinquefolia i Andromeda mariana 2 Asclepias obtusifolia i Clethra alnifolia 3 Epigaea repens 3 Eupatorium verbenaefolium 1 Euphorbia ipecacuanhae i Gaylussacia dumosa 2 frondosa o Ilex glabra o Kalmia angustifolia 3 latifolia 3 Myrica caroiinensis i Prunus serotina i Pyxidanthera barbulata 3 Quercus alba i prinus 3 " velutina 2 Solidago odora i Tephrosia virginiana 2 Vaccinium pennsyivanicum 3 I«I Generic Coefficient of the Pine-Barren Region We have called attention to the fact that Jaccard has determined the generic coefficient of the alpine Jura to be 54 per cent., and of the entire Swiss flora 26 per cent. For purposes of comparison, therefore, it is important to determine this coefficient for the pine-barren region of New Jersey. As we have ascertained, there are 555 species and 234 genera of plants native to this region. The generic coefficient is, therefore, 42.16 per cent., or in round figures 42 per cent., which is the proportion of genera to species in the pine-barren flora. A table will show how this percentage stands with respect to other regions.* Region New Jersey Pine-Barrens Miami, Florida Florida Keys Altamaha Grit Region of Georgia . . . Southeastern United States Region of Gray's Manual Central Rocky Mountains British Flora (Druce), including aliens Switzerland Species 555 796 533 797 6364 3413 2733 2964 2453 Genera 234 466 346 404 1494 821 649 734 659 Generic Coefficient Per Cent. 42 59 65 50 23 24 24 24 27 * Harshberger, J. W. Institute of Science, VII : li Vegetation of South Florida. , 1914- Transactions U'agner Free 1 82 VEGETATION OF THE NEW JERSEY PINE-BARRENS Families of Pine-Barren Plants It will be noted from the following two tables that the number of families of pine-barren plants is ninety-one. These families include 234 genera and 581 species of flowering plants. Of the 581 pine-barren species, 303 are known to have been collected at 6 localities and over; 41 1 have been gathered in from 3 to 6 localities, and 164 species in from i to 2 places in the pine-barrens. Ninety of the 164 have been reported from only one place, leaving 74 species collected in two localities. If we deduct 26 introduced species the total number of native pine-barren plants is reduced to 555 species. Six of the pine-barren species are of doubtful character or have been recently discovered, or rediscovered. Of the 91 families, 19 families, or 20.8 per cent., have only one species in this re- gion. With the exception of 27 families that comprise five or over five species, 64 are small, comprising not over four species to each family. We note the conspicuous absence* of such plant families as Equise- taceae, Selaginellaceae, Taxaceae, Chenopodiaceae, Portulacaceae, An- nonaceae, Papaveraceae, Cruciferae, Saxifragaceae, Platanaceae, Mal- vaceae, Polemoniaceae, Solanaceae, Bignoniaceae, Caprifoliaceae, Cu- curbitaceae, and Lobeliaceae. The most important families of pine- barren plants are noted in the table which follows. They are con- sidered important, because they contain the largest number of plant species and species which, from the synecologic view, are among the most important elements which enter into and render conspicuous the several plant formations and associations described previously. FAMILIES OF PINE-BARREN PLANTS Family Number OF Genera Number OF Species Family Number OF Genera Number OF Species 1. Ophioglossaceae .... 2. Osmundaceae 3. Schizeaceae 4. Polypodiaceae 5. Lycopodiaceae I I 2 6 I 2 2 2 9 4 13. Araceae 14. Xyridaceae 15. Eriocaulonaceae .... 16. Pontederiaceae 17 Juncaceae 5 I I I I 8 2 2 I 1 2 I 5 5 3 I 12 18. Melanthaceae 19. Liiiaceae 20. Convailariaceae .... 21. SmiLicaceae 22. Haemodoraceae .... 23. Amaryllidaceae 24. Dioscoreaceae 6. Pinaceae 3 S 9 3 1 1 1 I 22 1 1 I 2 3 2 71 76 8. Sparganiaceae 9. Najadaceae 5 I I I. Graminaceae 12. Cyperaceae I * Harper, R. M.: Review of Stone's Flora of Southern New Jersey, Torreya 12: 216- 225, September, 1912, especially page 220. VEGETATION OF THE NEW JERSEY PINE-BARRENS 183 FAMILIES OF PINE-BARREN PLANTS.— (Continued) Number OF Genera Number OF Species Family Number OF Genera Number OF Species 25. Iridaceae . . . 26. Orchidaceae. 27. Salicaceae 28. Myricaceae 29. Juglandaceae 30. Betulaceae 3 I . Fagaceae 32. Urticaceae 33. Loranthaceae 34. Polygonaceae 35. Caryophyllaceae. . . . 36. Nymphaeaceae 37. Magnoliaceae 38. Ranunculaceae 39. Lauraceae 40. Sarraceniaceae 41. Droseraceae 42. Iteaceae f43. Rosaceae 44. Pomaceae 45. Drupa- ceae . . . 46. Cae- salpini- Leguminosae \ aceae . . I 47- Papi- [ lionaceae Geraniaceae Linaceae Polygalaceae Euphorbiaceae Empetraceae Anacardiaceae Ilicaceae Aceraceae Vitaceae Rosaceae ■7 25 57. Hypericaceae. . . 58. Elatinaceae. . . . 59. Cistaceae 60. Violaceae 61. Cactaceae 62. Lythraceae 63. Melastomaceae. 64. Onagraceae. . . . 65. Haloragidaceae. 66. Araliaceae 67. Umbelliferae. . . 68. Cornaceae 69. Clethraceae. . . . 70. Pyrolaceae 71. Monotropaceae. 72. Ericaceae 73. Vacciniaceae. . . 74. Diapensiaceae. . 75. Primulaceae. . . . 76. Gentianaceae. . . 77. Menyanthaceae . 78. Apocynaceae. . . 79. Asclepiadaceae . 80. Convolvulaceae. 81. Cuscutaceae . . . 82. Boraginaceae. . . 83. Verbenaceae . . . 84. Labiatae 85. Scrophulariaceae 86. Lentibulariaceae 87. Rubiaceae 88. Caprifoliaceae. . 89. Campanulaceae. 90. Cichoriaceae . . . 91. Compositae .... Total 32 59 14 10 I 3 4 I 3 6 3 2 2 2 12 10 I I 6 15> 234 581 RELATIVE IMPORTANCE OF FAMILIES* Cyper.aceae 76 species. Graminaceae 71 Old Compositae 59 Old Ericaceae 32 Old Leguminosae 25 Orchidaceae 19 Old Rosaceae 17 Juncaceae 1 Onagraceae ^ 12 Labiatae J Lentibulariaceae 11 f^^^"^'^ > 10 species. Scrophulariaceae J '^ Polypodiaceae Melanthaceae Polygalaceae Hypericaceae Cistaceae Violaceae : . . Asclepiadaceae \- 6 Rubiaceae * Families in small capitals are important as to number of individuals. 1 84 VEGETATION OF THE NEW JERSEY PINE-BARRENS RELATIVE IMPORTANCE OF FAMIUES.— {Continued) 5 species. PiNACEAE Araceae Xyridaceae Smilacaceae Caryophyllaceae llicaceae , Lycopodiaceae Iridaceae Salicaceae Polygonaceae Linaceae Anacardiaceae Gentianaceae Campanulaceae Najadaceae Eriocaulonaceae Liliaceae Nymphaeaceae Droseraceae Euphorbiaceae Lythraceae Melastomaceae Haloragidaceae Umbelliferae Primulaceae Apocynaceae Convolvulaceae Ophioglossaceae Osmundaceae Schizaeaceae Sparganiaceae * Introduced or relatively not abundant. Alismaceae Convallariaceae. Amaryllidaceae. . Salicaceae Myricaceae. . . . Jugiandaceae. . . Betulaceae Vitaceae Araliaceae Boraginaceae . . . Verbenaceae. . . . Caprifoliaceae. . . Typhaceae *. . . . Pontederiaceae. . Haemodoraceae . Dioscoreaceae . . . Loranthaceae. . . Santalaceae Magnoliaceae . Ranunculaceae *. Lauraceae Sarraceniaceae . . Iteaceae Geraniaceae *. . . Empetraceae . . Aceraceae Elatinaceae Cactaceae * Cornaceae .... Diapensiaceae. Menyanthaceae. 2 species. CHAPTER XIV BIOLOGIC TYPES OF PINE-BARREN PLANTS Raunkiaer* divides plants according to the position of the winter-buds with reference to the soil surface. The winter-buds of the phanerophytes are borne at least 30 centimeters above the surface of the soil. Those of the megaphanerophytes are borne 30 meters above the earth; those of the mesophanerophytes, 8 to 30 meters; microphanerophytes, 2 to 8 meters; nanophanerophytes, 0.3 to 2 meters. The chamaephytes are those plants whose winter buds are borne immediately above the sur- face of the soil. The hemicryptophytes have winter-buds just below the earth's surface in the upper crust of the soil. The cryptophyies bear buds some depth below. They are divided into the geophytes, which have buds under the surface of the soil. The helophytes and hydrophytes have their buds entirely covered with water. Therophytes are the annual plants that reproduce by seeds. Martin Vahlf proposed the terms dia- geic and epigeic. The diageic plants are geophytes that grow trailing under the ground, while epigaeic plants are phanerophytes and chamae- phytes that develop mainly above the surface of the soil. An attempt is made here to group the native species of the pine-barren region of New Jersey under the above growth forms, but this has not been easy, because our knowledge of many species, either described in the manuals or as herbarium specimens, is very incomplete and unsatis- factory. It is, therefore, probable that mistakes have been made. Some no doubt will neutralize others, but such errors of determination are unavoidable when the descriptions and the specimens preserved in herbaria do not enable the statistician to reach determinative con- clusions. The percentage of different growth forms in a region arranged in a series, or gamut, gives a picture or spectrum of the vegetation in the words of Raunkiaer: " Ich werde deshalb der Ktirzes halben in folgenden * Raunkiaer: Types biologiques pour la geographic botanique. Bulletin de I'Aca- demie des Sciences du Danemark, 1905. t Vahl, Martin: Les Types biologiques dans quelques Formations Vegetaies de la Scandinavie, Acad. Roy. des Sci. et des Lettres de Danemark, 191 1 : 322-323; The Growth Forms of Some Plant Formations of Swedish Lapland. Dansk. Bot. Arkiv., i, No. 2, 1913. 185 1 86 VEGETATION OF THE NEW JERSEY PINE-BARRENS eine solche statistich-biologische Uebersicht als Spektrum bezeichnen, als biologisches Spektrum oder Pflanzenklimaspektrum." He has worked out on theoretic grounds a normal spectrum, which is given in the adjoining table, where S = stem succulents; E = epiphytes; Ph = phane- rophytes; Ch = chamaephytes; H = hemicryptophytes; G = geophytes; HH=helophytes and Th = therophytes. Region Pine-Barrens of New Jersey Miami Florida Region Florida Keys St. Thomas and St. John . . Seychelles Aden Transcaspian Lowlands * . . Pamir * Death Valley Samos Libyan Desert Cyrenaica Normal Spectrum Number 458 793 527 904 258 176 768 5'4 294 400 194 375 400 Percentage of Species in Each Growth Form ■23 14 22 32 5.23,30 10, 23, 24 -. 7, 26 I I I 26 9 12 9 6, 17, 20 Ch H 10 16 20 12 6 27 7 12 7 13 21 ■4 9 38 28 ■9 9 12 19 27 63 18 32 20 19 27 HH Th 18 20 •4 16 '7 41 •4 42 33 42 50 >3 Under Ph for St. Thomas and St. John islands, for the Seychelles, Aden and the Normal Spectrum, the percentages are arranged as MM, M and N, i. e., Megaphanerophytes, Microphanerophytes and Nano- phanerophytes. It will be noted that in the flora of the pine-barrens the hemicryptophytes are the most abundant, followed by the helo- phytes, the phanerophytes, the therophytes and the chamaephytes. Without adopting this classification of Raunkiaer for the arrangement of plants into biologic types I will use instead the classification of plants into the following types: trees, small trees, shrubs, undershrubs, peren- nial herbs, annual herbs and into evergreen and deciduous-leaved species. Pine-Barren Formation Pinus echinata. " rigida. Juniperus virginiana. Quercus alba. coccinea. Trees Quercus marylandica. " prinus. stellata. triloba ( = falcata). Sassafras variifoiium. * Paulsen, Ove: Studies on the Vegetation of the Transcaspian Lowlands, 1912: 135- '73- VEGETATION OF THE NEW JERSEY PINE-BARRENS 187 Myrica carolinensis. Quercus ilicifolia. prinoides. Crataegus tomentosa ( = uniflora). Clethra alnifolia. Andromeda (Pieris) mariana. Arctostaphylos uva-ursi. Comptonia asplenifolia. Corema Conradii. Gaylussacia resinosa. Hudsonia ericoides. Aristida dichotoma. Agrostis hyemalis. * Gyrostachys Beckii. Linum medium. Crotonopsis linearis. Polygonella articuiata. * Bartonia panicuiata. Shrubs Dendrium buxifolium. Gaylussacia dumosa. frondosa. Kalmia latifolia. Undershrubs Hudsonia tomentosa. Kalmia angustifolia. Smilax glauca. Vaccinium pennsylvanicum. " vacillans. Annual Herbs * Bartonia virginica. * Dasystoma pedicularia. Gerardia Holmiana. Melampyrum lineare. Trichostema dichotoma. lineare. Gnaphalium purpureum. Annual, or Biennial. Cuscuta arvensis. Colorless Parasitic Plants Cuscuta compacta. Pteris (Pteridium) aquilinum. Paspalum setaceum. Aristida purpurascens. Pe Pan cum depauperatum. Lindheimeri. meridionale. oricola. villosissimum. Commonsianum. Addisonii. tsugetorum. columbianum. thinium. sphaerocarpon. Ashei. Amphicarpon amphicarpon. Stipa avenacea. Cyperus cylindricus. Grayi. filiculmis macilentus. Scleria pauciflora. Carex pennsylvanica. Xerophyllum asphodeloides. Cypripedium acaule. Comandra umbellata. Aletris farinosa. Arenaria caroliniana. Baptisia tinctoria. Tephrosia (Cracca) \irginiana. Stylosanthes biflora. Ferns Woodwardia areolata. RENNiAL Herbs Meibomia rigida. obtusa. Lespedeza frutescens. hirta. " angustifolia. Galactia regularis. Euphorbia ipecacuanhae. Ascyrum hypericoides. Helianthemum canadense. Lechea minor. racemulosa. villosa. Kneiffia linearis. Pyrola americana. Epigaea repens. Gaultheria procumbens. Pyxidanthera barbulata. Gentiana porphyrio. Asclepias tuberosa. amplexicaulis ( = obtusifolia). Breweria Pickeringii. Epilobium angustifolium. Schwalbea americana. Monarda punctata. Lycopus sessilifolius. Mitchella repens. Hieracium venosum. Gronovii. Lacinaria graminifolia pilosa. VEGETATION OF THE NEW JERSEY PINE-BARRENS Pinus rigida. " echinata. Juniperus virginiana. Xerophyllum asphodeloides. Cypripedium acaule. Arenaria caroliniana. Hudsonia ericoides. tomentosa. Ilex glabra. Evergreen Species Corema Conradii. Arctostaphylos uva-ursi. Dendrium buxifolium. Epigaea repens. Gaultheria procumbens. Kalmia latifolia. Pyrola americana. Pyxidanthera barbulata. Mitchella repens. Woody Deciduous-leaved Species Comptonia asplenifolia. Myrica carolinensis. Quercus alba. " coccinea. " ilicifolia. " marylandica. " prinoides. " prinus. " stellata. triloba ( = falcata). lnconsDicuou«. Apetalous. Sassafras variifolium. Crataegus tomentosa ( = uniflora). Andromeda (Pieris) mariana. Clethra alnifolia. Gaylussacia dumosa. frondosa. resinosa. Vaccinium pennsylvanicum. " vacillans. Creanish While Whita. Pink or Red. Figure 155 Tabulation of flower colors of pine-barren plants. Colors of Pine-Barren Flowers* As the colors of flowers are important in describing the vegetation of a country, the following analysis of the plants of the pine-barren forma- tion proper is presented. The grasses and sedges are not included. Yellow 29 White 26 Pink or Red 10 Purple 16 Blue or Violet 9 Variegated 4 It will be noted in the above enumeration that yellow and white are the most prevalent flower colors in the pine-barren formation. The * Allen, Grant: The Colours of Flowers, 1882. VEGETATION OF THE NEW JERSEY PINE-BARRENS 189 grasses and sedges are excluded, for although the protuberant anthers of the grass and sedge flowers give color to the vegetation, yet it can hardly be said that such plants have flower colors. For the same reason the pines and oaks have been omitted from consideration. If we take the two less specialized colors, we find that there are 55 pine-barren species with yellow and white flowers. Taking the more highly specialized flower colors, there are 39 species of pine-barren plants with pink, red, purple, blue, violet and variegated flowers, so that the unspecialized flower colors exceed the specialized by some 16 species. White-Cedar Swamp Formation In arranging the species, according to biologic characteristics and growth forms, no reference will be made to the synecology of the white-cedar swamp formation. The association of species has been described in a former section. Trees Evergreen: Deciduous: Chamaecyparis thyoides. Acer carolinianum. Magnolia glauca ( = virginiana). Nyssa sylvatica. Kalmia latifolia. Rhododendron maximum. Shrubs Alnus serrulata ( = rugosa). Gaylussacia frondosa. Myrica carolinensis. Vaccinium atrococcum. Uicioides mucronata. corymbosum. Rhus vernix. " macrocarpon. Azalea viscosa. Ferns Dryopteris simulata. Osmunda cinnamomea. Lycopodium alopecuroides. regalis. carolinianum. Schizaea pusilla. Chapmanni. Grasses Calamovilfa brevipilis. Panicum lucidum. Danthonia exilis. spretum. Panicularia obtusa. Sporoboius serotinus. Panicum ensifolium. Torreyanus. Sedges Carex bullata. Eriophorum tenellum. canescens disjuncta. virginicum. Collinsii. Rhynchospora alba. exilis. axillaris. interior capiilacea. fusca. livida. glomerata. oblita. " gracilenta. trisperma. oligantha. Walteriana. Torreyana. Cladium mariscoides. Scieria triglomerata. Insect-catching Plants Drosera longifolia. Utricularia gibba. rotundifolia. intermedia. Sarracenia purpurea. juncea. Utricularia cornuta. igo VEGETATION OF THE NEW JERSEY PINE-BARRENS Orchids Arethusa bulbosa. Gymnadeniopsis clavellata. Blephariglottis blephariglottis. Limodorum tuberosum, ciliaris. Pogonia divaricata. cristata. " ophioglossoides. Gymnadeniopsis integra. Aquatic Plants Castalia (Nymphaea) odorata. Orontium aquaticum. Parasitic Phoradendron flavescens. Perennial Herbs Spathyema foetida. Decodon verticillatus. Xyris Congdoni. Rhexia virginica. Eriocaulon decangulare. Sabatia lanceoiata. Juncus aristulatus. Asclepias rubra. caesariensis. Gerardia racemuiosa. " pelocarpus Lycopus sessilifolius. Abama americana. Aster nemoralis. Tofieldia racemosa. Bidens trichosperma tenuiloba. Helonias buliata. Eupatorium leucolepis. Lophioia aurea. " resinosum. Hypericum boreaie. Sclerolepis uniflora. Viola lanceoiata. Solidago neglecta. Flower Colors of White-Cedar Swamp Plants If we tabulate the colors of the flowers of the white-cedar swamp plants, we find that there are no blue flowers and no variegated flowers among the true cedar swamp plants. Excluding the white-cedar, the grasses and the sedges from consideration, the enumeration stands as follows: Yellow 14 Purple 8 White 21 Brownish-green 3 Pink, or red 9 The sum (35) of the yellow and white flowers exceeds the sum (20) of the pink, red, purple, and brown colors by 15. Biologic Types Found in Marsh Formations Under this category may be enumerated the plants native to the tree swamps and open marshes of the pine-barren region. The important biologic groups of plants are given with reference to their growth forms,* as they impress the general observer of the native vegetation. Ferns Osmunda cinnamomea. Woodwardia virginica. " regalis. Grasses Calamagrostis cinnoides. Panicularia obtusa. Panicularia canadensis. " pallida, nervata. ■ * In connection with growth forms consult Drude, O.: Oekologie der Pflanzen. VEGETATION OF THE NEW JERSEY PINE-BARRENS 191 Carex Barratti. bullata. folliculata. intumescens. lurida. Cyperus strigosus. Perennial Her Xyris Congdoni. fimbriata. " torta. Eriocaulon compressum. decangulare. Juncus acuminatus. " aristulatus. Apios tuberosa. Dioscorea villosa. Phoradendron flavescens. Polygonum Careyi. Helonias bullata. Lilium superbum. Iris prismatica. Gymnadeniopsis integra. Polygonum emersum. Linum striatum (biennial). Polygala brevifolia. Nuttallii. Viola lanceolata. " primulaefolia. Hypericum virgatum ovalifolium. Rhexia mariana. virginica. Ludvigia alternifolia. hirtella. Sedges Dulichium arundinaceum. Eleocharis tricostata. Rhynchospora gracilenta. macrostachys. Scirpus americanus. Scleria minor. iS WITH Sedge- or Grass-like Leaves Juncus canadensis. Juncus effusus. Zygadenus leimanthoides. Melanthium virginicum. Sisyrinchium atlanticum. Gyrostachys praecox. Climbing Plants Smilax W'alteri. Parasitic Plant .Annual Herbs Rotala ramosior. Perennial Herbs Ludvigia linearis. sphaerocarpa. Proserpinaca pectinata. Oxypolis rigidior. rigidior longifolia. Lysimachia terrestris. Sabatia lanceolata. Utricularia cornuta. Coreopsis rosea. Soiidago fistulosa. neglecta. uniligulata. Doellingeria humilis. umbellata. Helianthus angustifolius. Aronia arbutifolia. nigra. Decodon verticillatus. Woody Plants (Undershrubs, etc.) Rubus hispidus. Spiraea latifolia. Capsular Fruits: .'\zalea viscosa. Chamaedaphne calyculata. Clethra alnifolia. Itea virginica. Kalmia latifolia. Leucothoe racemosa. Xolisma ligustrina. Baccate Fruits: Ilex glabra. Shrubs Rhus vernix. V'accinium atrococcum. " corymbosum. Drupaceous Fruits: Ilex laevigata. " verticillata. llicioides mucronata. 192 VEGETATION OF THE NEW JERSEY PINE-BARRENS Evergreen: Chamaecyparis thyoides. Pinus rigida. Deciduous: Drupaceous Fruits: Nyssa sylvatica. Sassafras variifolium. Trees Pomaceous Fruit: Amelanchier intermedia. IVinged Fruit: Acer rubrum. Betula populifolia. Capsular Fruit: Liquidambar styraciflua. Aggregate Fruit: Magnolia glauca ( = virginiana). Flower Colors of Marsh Plants The colors of the flowers of the marsh plants of New Jersey present a greater variety than do either the pineland or the white-cedar swamp plants. Enumerated, the colors are as follows: Inconspicuous i No petals i Greenish 2 Yellow 22 Greenish-white 1 White 20 Pink or red 7 Purple 4 Blue 2 Greenish-brown 3 Yellow and white, as before, are the prevailing tints of flowers among marsh plants. Tabulating the flower colors for the pineland, white-cedar swamps, and marshes we have (Fig. 155): Pineland White-Cedar Swamp Marshland Total Inconspicuous. . No petals Greenish Yellow Greenish-white. White Pink or red . . . Purple Blue Greenish-brown Variegated 29 26 10 16 9 14 21 9 I I 2 22 I 20 7 4 2 3 I I 2 65 I 67 26 28 1 1 6 4 Consulting the table and the figure (Fig. 155) we discover that white (67), yellow 65, purple (28), pink or red (26), and blue (11) are the most frequent flower colors in the pine-barren region of New Jersey and in the order named. The other colors are unimportant. CHAPTER XV PHYTOPHENOLOGY OF PINE-BARREN VEGETATION That phase of botanic science which concerns itself with a study of the course of vegetation referred to the seasons is known as phytophenom- enology, shortened to phytophenology. It is of interest to consider the period of plant activity and the period of dormancy induced by the approach of the winter cold. The bursting of the leaf and flower buds, the unfolding of the first leaf, and its final fall in the autumn are all matters of ecologic interest. The floral procession of an entire year has been studied in only a few instances, and the following account is one of the few descriptions of the procession of plant development and plant flowering, from the phytogeographic standpoint, that is, considered from the standpoint of the whole vegetation of a definite region. The re- corded observations in America, heretofore, have dealt in the main with individual plants, such as the unfolding of the first leaf, the opening of the first flower, the withering of the last blossom, the fall of the leaf and the development of the fruit. These results have been tabulated for chosen species, but only a few consistent attempts have been made to study the procession of plants from early spring to late fall from the regional aspect, and few attempts have been made to make deductions based upon such a study. The following tabulation of 548 pine-barren species, as far as the flow- ering and fruiting periods are concerned, is based on the field and herb- arium notes of the writer and upon the work of Bayard Long, who has described succinctly his method of gathering the data in Stone's " Plants of Southern New Jersey," where the facts are recorded under each plant species. The construction of the graphic table has occupied some time. The table shows the flowering and fruiting periods of 548 pine-barren plants, seven plants less than the total number of native species, from the first of March to the end of November, a period of nine months, leaving only three months as the time when the pine-barren vegetation, taken as a whole, is entirely dormant. This period can be shortened to seven months by excluding the months of March and November, for only about four species show any activity in March and none certainly in November. It is shortened to six months if the climatic records are 13 '93 194 VEGETATION OF THE NEW JERSEY PINE-BARRENS considered. Presumably the growing season of pine-barren plants is between the last killing frost of spring and the first killing frost of au- tumn. At Vineland, according to the records of the U. S. Weather Bureau, the average date of the last killing frost is April 19, and the latest recorded date May 13. The average date of the first killing frost in autumn is October 19, the earliest recorded date October 2. If we take the average dates the season of growth is exactly six months, or one hundred and eighty-three days in length. This statement was written in 1912. Since then, three years later, in January, 1915, there has ap- peared, while this monograph was being recopied, a work by Taylor* in which this matter is discussed with reference to the region within 100 miles of New York City. On the map, Plate 5, of Taylor's memoir, the region north of the heavy black line, which runs north of the northern part of New Jersey, has a growing season of one hundred and fifty-three days, or less, while the country south of this line has a growing season of one hundred and sixty-four days, or more, and in this part of the map the New Jersey pine-barrens fall with one hundred and fifty-nine to one hundred and eighty-two days as the length of the growing season shown on the face of the map, a close approximation to one hundred and eighty-three days, the average length of the growing season deter- mined above from the Vineland records. This determination of the length of the growing season, as of importance in the geographic distribu- tion of plants, is in line with the labors of Cleveland Abbe, who has pub- lished an important monograph on the subject. f To come back to our original argument, seven months from the be- ginning of April to the end of October represent the grand period of activity of pine-barren vegetation, leaving five months when the vege- tation, as a whole, is practically dormant. Consulting the tables, it will be seen, too, that April is a month in which very little activity is shown, so that if we exclude the early spring plants (and there are about 39 such plants, or 7 per cent, of the whole), the grand period of vegeta- tion is shortened to six months. The principal flowering and fruiting periods are not reached until July by the majority of pine-barren plants, and the active period of flower and fruit production for this majority is crowded into the three months of July, August and September. The cul- * Taylor, Norman: Flora of the Vicinity of New York. Memoirs New York Botani- cal Garden, v: 35-36, map, Plate V. t Abbe. Cleveland: A First Report on the Relations between Climates and Crops. Bull. 36, U. S. Dept. of Agriculture, Weather Bureau, 1905, pages i to 386; consult also an important paper by William G. Reed and Howard R. Tolley: Weather as a Business Risk in Farming. The Geographical Review, 11: 48-53, July, 1916. The risk has been graphically and mathematically presented. VEGETATION OF THE NEW JERSEY PINE-BARRENS I95 mination of this phenomenon takes place in August. So much for the general features of the phenologic table. It should be mentioned that the several months are divided in the phenologic table into six periods of five days each, and this division of the months is sufficiently accurate for phenologic purposes, if we make each of the growing months on an average thirty days long. The flowering periods are indicated in the table by full black lines and the fruiting periods by dotted lines. It will be noted that some flowering periods are short and others are long. Some fruiting periods overlap the period of flower production, but in most cases the production of fruit follows the flower period and is separated from it by an interval of time. From a visit to the pine-barren region early in November, when every plant had entered the resting condition, it may be said that October ends the period of growth. But October is a month when the autumn plants are preparing for the period of dormancy and when very few plants flower except some fall Compositae. it may be considered to be a month of preparation rather than a month of active growth. Very few new phenologic events are begun in October. They are begun in other months and are finished in October. If this view of the month's activities is considered, it will be conceded that we could exclude Octo- ber from the months of most active growth, leaving only five months out of the twelve when the pine-barren plants are in an active stage of development. These facts are of importance to a man engaged in active agricultural operations in the region. Data from my field notes corroborate the facts displayed in the fore- going table. Excluding the smaller herbaceous plants which were found in flower and a few flowering shrubs, the pine forest on April 20, 1910, was in its winter aspect. Ten days later, on April 30, 1910, the vegeta- tion had undergone a change. It was noted on that date that Quercus prinus was half in leaf. Quercus marylandica and Quercus ilicifolia displayed their male catkins and female flowers and were in young leaf- age. The leaves of Quercus stellata and Q. tinctoria were small. The leaves of the red-maple, Acer rubrum, were expanded almost entirely, but the sour-gum, Nyssa sylvatica, and Magnolia glauca were still bare of leaves with their buds unopened. The new shoots of Pinus rigida on May 4, 1 910, had reached a length of four inches, and on this date the young leaves of Liquidambar styraciflua and Nyssa sylvatica were seen. By June i all of the above trees were in full foliage, but the leaves re- tained their juvenile light green color, contrasting with the dark greens of the pitch-pine, Pinus rigida. It will be seen from these data that the month of May is practically the one in which the leafage of the deciduous 196 VEGETATION OF THE NEW JERSEY PINE-BARRENS Flowering and Fruiting Periods of Pine-Barren Plants Heavy lines, flowering periods; dotted lines, fruiting periods Name of Plant I Botryohlum abliquum tT 2 Botryohium obllquuir dieeectt. ' 3 Oamunda regilie "■+""" 4 Oeir.unda cin lairomea 5 Sohizaea puallla 6 Lygodiun palmatuD 7 Pteridlua aiullinum 8 Woodwardla virginioa 9 Woodwardla areolata 1 "^ 10 Aaplenlum platyneuron t H Aaplenlum filu-foenilna 13 Polyetiohum aoroetlcholdea ■f :::: 13 DryopterlB thelypterla .IE .», 14 Dryopteria Blmulata 15 Phegopterls dryopteria 16 Lyeopodium chapDanll 17 LyoopcdluB alopeouroldee ...:::::;":: 16 Lyeopodium caroHnlanum 19 lyeopodium obaourum 20 Pinua vlrglalana 21 PlEua eehinita 23 Pinua rlgldj 33 Chamaeoyparis thyoldes .... 34 Juniperua Tlrginlana 25 Typha latlfolla """ 06 Sparganium aiterioanuir. ;;;1::;:;" 27 Sparganlua amerlcanum andrc 36 FotsEogeton oakealanua 29 " eplhydrue 30 " oonfervoldea k:z3:^ ;!!!i;'"'"Sr":n 32 Sagittaria graitlnea 33 KrlanthUB aa-joharoldee 34 Andropogon aooparlua 35 " coryirbooua abbrevlatu ttI 35 " virglnloua tt 1 37 Sorghaatrum nutana 1 ' 36 Paepalum paarrTophiluit 39 ' r-iJbeacens irrr 1 r jTit 40 " muhlenbergll _r III' 41 " eetaceuir, 43 AEPhlcarpon arrpnioarpon 1 1 lllllllll VEGETATION OF THE NEW JERSEY PINE-BARRENS 197 Flowering and Fruiting Periods of Pine-Barren Plants 43 ?ynt^.e^iBI^a filifgrrii 44 Paniour verrucoaum 45 " phlladelFhlourr. 46 " viriTAt-un 47 " " cuberae 48 " af;i-03toijro 49 " Icnslfolirr,! f,0 " dop^upcratum 51, " dichotonuir. b3rbul3tutTi lucidura Glutei cpret'jn ' llndheineri louc&thrix ' inpridicnale ' criccla ' vil]oji93rnum ' fBeuJspubeecer.B ccniTiO.'iBianum " addiDonil " tsugeto: " colunoianuir » " thlriur. " en^iloliun " apicrocarpon " ashel " cl'igoafvnthee " eoop3rii.r. 72 " cryptanth'jm 72 " ocabriuGculurr. 74 Cenohrue caroliniarus 75 Aristida dichotora 76 " jraolliB 77 " FurP'JraDr;era 7ti Stipa jrepac^a 79 Vuhlerberjla caplllarl SO Spcrobol»;8 torreyar.uB 81 " Bcrotirua fl2 AgroBtis marltJma 198 VEGETATION OF THE NEW JERSEY PINE-BARRENS Flowering and Fruiting Periods of Pine-Barren Plants Rare of Plant ■M ..,11-. J » tfnibffi ■?atp&e 65 OalarMErostla olnnoldeo 36 Calartvilfa brevlpllle S7 Deschampaia flexuoaa 68 Dantnonla aplcata 89 ' aerloea 90 " epUU 31 GyE-.opogon iitblguua "iS Triplasla purpurea 93 Kragrcetls plloaa '94 " pectlnicea 55 Panicularla canadenaia 96 " obtuaa 97 " nervata <:>8 " pallida 09 ' aoutlflora 'IOC Fejtuoa oetoflora 101 Cyperua flaveaoene 103 ' dentatua 103 " BtrlgoauB 104 " hyatriolnue 105 " fetrofractus 105 " cyllndrloua 107 ' grayl 108 "filioulnls macllentus 109 Dullshlum aiundlnaseum 110 Eleooharls robblnaii 111 " cllvaoeae 113 ' obtuaa 113 " aoioularls 114 " tuberculosa 115 " tcrreyar.a 115 " tncostata 117 " tenuis il8 Stenophyllua oaplllaris 119 Fimbiiatylis autumnalls 120 " oaatanea 121 Sclrpus aubteriLlnalls 1S3 ' anerloanua 123 " llneatus 124 " lonjll 125 " cyperlnus J26 ' erlophorum :• VEGETATION OF THE NEW JERSEY PINE-BARRENS 199 Flowering and Fruiting Periods of Pine-Barren Plants Name of Plant 'i'T"i°! iT^r^i'' 137 128 189 130 131 133 133 134 135 136 137 138 139 140 141 143 143 144 145 146 147 148 149 150 151 153 153 154 155 155 157 160 161 163 163 a 64 165 166 167 168 Erlophorum tenellum " TirglnlouB Rynahosrota nacrostachya " maocoatiohya Inundat^ " graollenta " oltgantha " alba " ralllda " knieeVerr.li " gloir.erata " jlor3rata leptooarpa " BEallU " fllifolia " ajclllarie " fusca " tcrreyana Cladlum rrarleooldea Sclerla trljlOKerata " ii.Uor ■ reticul iria torreyana " pauolflora Carex col Una 1 1 " folliculata " Intuneacena " lupullna " bullati " lurlda " lacuatrle " veatlti ' walterlara " tarrattll " crlnlta " ewanil ' trloera " obllta " llvlda ' peinsylvanlca " unbellata '.onsa " enlllB • arrecter.a ■ tTuhlenfc^^rgll ■ Interior cjpiuacea 200 VEGETATION OF THE NEW JERSEY PINE-BARRENS Flowering and Fruiting Periods of Pine-Barren Plants Nace of Plant July I . Aujuat Beptembei getobfr 168 Carex atlantloa I'O " caneeoens dlejunota I'l " trieperira 1'3 " eooparla 1"3 " alboluteecena l'* Arieaeira puelllun I'S Peltandra vlrglnloa 1'6 Orontlum acjuatloum 1" Aoorue calamue 1'8 Xyrlo torta 179 » congdonl 180 " carollnlana 161 " finbriata 163 " arenicola 183 ITlocaulon aeptmgulire 184 " conipreaeuDi 185 " decingulare 186 Pontederia oordata 187 JuncuB effuBue 133 " teruis 18S " dichotoirua 190 " marglnatUB ISl " arlatulatuB 193 " pelocarpuB 193 " BllUarle 194 " caesarlenais 195 " BolrpoideB 196 " canadenala 197 " aouitilnatuB 198 " deblllB 199 Tofleldla racemoBa. 3CC Abama ar.erlcana 2C1 Xerophylluiii aophodeloldea ?03 Helonlas buUata 203 Chroaperma niufloaetoxicum 204 ZIgadenuB lelmantholdee 205 Uelanthlum yirglnlciim 305 Uvularla eeBBlllfolla 207 " nitlda 208.LillujD auperbuB 209 Al«trlB farlnoBa 210 Polygdnatum comButatum VEGETATION OF THE NEW JERSEY PINE-BARRENS 201 Flowering and Fruiting Periods of Pine-Barren Plants Naie of Plant £11 Uedecla vlrglnlana 312 Sii.lla» tair.iruUa ;i3 " rotundlfolla 214 " gljuoa 215 " laurifJlla 216 " VJalteri 217 Gyrotheoa tinstorla 216 Ryroxis hir-ita 219 Lophlola aurea 220 Diosoorea viUcrr. 2i\ Irie prisiBtica 222 Sisyrinphiuir rucronatum 22S " atlantlcum 224 Cyprlpedium aoaule 225 GymnadenitpciB Integra 226 " clavellata 227 Blepharlglottla orlstata :26 " cillaris 22S " blepharigJottia 23C Pogenla cphioglosBC/idee i31 " dlvarlcita 232 iBctrla verticUlata 233 Arethuea bulboea 234 Liiiodoruir. tuberoouoj 235 CyroBtaohys cernua 233 " praeoox E3T " vernalle 336 " becLii 639 Liatera auatralie 24C PeranluLi pubescpna 241 Lertcro''.lB loeaelii 342 PopuluB jrar.dldertati 243 Salix .iljra 344 " hur-.iUe 245 " triBtia 246 Myiioa carolinensle 247 Comptcnia aapler.lf oils 34B Hicoria glabra 249 Betula poiJulifolla 350 Alr.ua rugoaa 351 OuerouB ooocinea ir. 3 17 - fl srldar.uir. 313 " 3t liatun 319 Pol •gaLi lutca 320 " cruciita 321 " brevlfolia 32? - v»rticilla',a 333 - ai'biJua 324 • virideaccns 325 • nariar.a 723 " r.uttallil 3? 7 - poly^aira 3c£ Crctcr.cpsia lir.eariB 329 Aoalypha .'racilena 33C F'jphorbia iceoao'j-.rhae 331 Cores^a cenradii 333 Rhus copal lir.ur 333 " verr.ix 334 " toxicodendron 335 Ilex cpaca ^25 " ilura 112^ 9 f.r.j.rj'.P'i' ?. 204 VEGETATION OF THE NEW JERSEY PINE-BARRENS Flowering and Fruiting Periods of Pine-Barren Plants Kaite cf Plant Marrh ^^'"-;-| V^y -T"f" -T-iy .„.,, 337 Ilex vertlolllata Li... ...... 339 Illoioidee r.ucronata ■■■■ ••••• ".•••Ml 340 Acer rubrum caroUnianunj ""' •••» ' 343 AecyruB etano 343 " hypericoidee " "T"' , "♦••• \" I /"';'^'' 34S Sarothra gentlanoldee " it T'V"^' .1..— 1j«.1 . ' k 353 Hudsonia tomertoBa ' ■■■ >>»►<■«« 353 " erlcoidee '■■■■ ^tioiaaj 354 Leohea ir.lnor 355 " raceiLUlcoa * 356 " vllloea »• > • » 357 " Earltlma 356 ' legjettil '•••»! • t • « 361 " flrtbrlatula ■■^■. 364 " lanceolata ' 366 notala lacoaior | >••••! • •••w. >••«•, ^^ • ». :::::::::::::: 1 ■ >■> 2::: — ?:::: > .»»••• , 1 '< > 1 • • • • ^;j»» Fffl-H i VEGETATION OF THE NEW JERSEY PINE-BARRENS 205 Flowering and Fruiting Periods of Fine-Barren Plants 3''3 X/ioiffia llne«ci9 3£0 * IcngipfJleellata 381 " pumlla 383 Prooerpinica pcctinata 333 HyriOFhyll-iE hunlle :e4 Aralla audieaulle 365 CxypoiiB rieldlCr 155 OtypsUe riilditr longifc 337 Xyaaa eylvatlca 3Q8 Clethr^ alnl folia 369 Pyrola aaerlcina 390 • elliptlca 091 • chlorar.tta 392 Chlcapblla tacula'.a 393 • unb'llata 334 aonotrepa «r.lflera 335 Sypopitys hypopithya ;£5 Azalea rjJ.riora "a' " viasota 396 ' • glauca 399 RhO'loienlron maxiawr. 400 Dendrl'io buxifoilui, 401 Xalula ar.£ustirolxa i02 ' latlfclla 403 Leucothoe laccmoaa lot Pierls irarlar.a 4C5 XcliBia lljustrlr.a ♦03 rplgaea lepena 407 C!iaL-.aeclaphne ealyculitd 408 "iul'.herla prsiucber.a 409 Aretoatapbyloi ura-jrsi 410 Cayluasasla fronlcaa 411 * ilunoaa 412 ■ bacceta 413 Vacslniuin ncryobosuiL 414 viriatum 415 caeeirlcnic 416 atroooocuc 417 rer.n9ylvir.ic 418 vaeslllana 419 Oxyccccua itacrooarpua 430 Pyxldanthera barbulata 521 Lyaltrachla ciuadrlfCTila 206 VEGETATION- OF THE NEW JERSEY PINE-BARRENS Flowering and Fruiting Periods of Pine-Barren Plants !'an-.e of Plant 423 Lyeinachla^ terrsatrie 433 TrieTitalis borealia 434 Sabatia lanoeolata 435 Oer.tiana porphyrio 436 Bartonia paniculata 427 " virslnioa 428 Liir.nanther.UK lacuncaua 4"9 ApccynuiB androaaeirifcli 430 " iLediujn 431 " cannabinum pubeacenB 433 Aecleplaa tuberoea 133 " rubra 434 " aicrlexicaulie 435 " varlegata 436 " syrica 437 " vertioillata 438 Brewerla piokertngii 43S Ipomoea pandurata 440 Convolvulus seplum 441 Cuecuta arvensla 443 " coF.paota 44: liyosotls laxa 444 OnoBffiOdluin virginidr.um 445 Verbena urtioifolia 145 " anguBtlfolia 44' TrlohoBtema dichotorrum ',48 Scutellaria integrlfbll.i 44S. Etdchys hyeeoplfolia ■iSC Mcnarla punctata 151 Hedeona pulegicidea 152 Cilnci'Oiiuir. vjlgare 453 Kcelila vertioillata 454 " mcana 455 " cutlca 455 Lyoopja virgirl-u8 457 " seieilifolius 458 Llnarla car.aderalB 45S Oratiola aure» 4S0 Veronica peregrina 461 DasyatoKa pcdlcularia 493 • flava 463 • Tjfglnloa VEGETATION OF THE NEW JERSEY PINE-BARRENS 207 Flowering and Fruiting Periods of Pine-Barren Plants Kane of Plant 454 Oerardia raceiruloea 4£5 •* holmiana •IGG Schwilbci amerlcana 43*? UelappyruB lineire 468 Utricularla cornuta 169 " juncea I'C • virgatula I'l • fibroaa ♦»2 • subulata 475 • clclstcgir.a 4»1 • clandeatlr.a <•»» • glbba ♦'S • Intermedia «■" • mflata ♦ T8 ' piirpurea 4^ Olvtnlandia unifldra ♦ttO Cephalanthu» occider.tsilli tei mtehella repeoe 4 82 Dlodi4 ter** 4 93 Galium plloatia 1C1 " • p-jnottiuloouii 455 Viburnum caaonoldpe 11 86 " r.uiun HE" Speeuliria r"follat* 1488 Lobelia inflata 169 * nuttallll -L... 532 " neicaralls z I ... 533 " denoauB M t; 534 Eosllingeria mbellata hue ilia 1 ....£ 536 Baochario hallrrifolia •••••>•• 537 Antennarla neglecta [ ■ T T 538 " plantaglnlfolla 539 » parllnil 540 Anaphalla margarltaoea |541 Gnarhallum obtuai folium i 543 " purpureum T 544 » dlrarlcatua 545 Coreopala rsaea 546 Bllsns conrata T 547 " tricliOaperma tenulloSa ^1f uPneelo tomentoaua Hill VEGETATION OF THE NEW JERSEY PINE-BARRENS 209 trees takes place, and October is the month in which the death of the foHage leaves of such trees occurs. The period of active metabolism of mature leaves is about four months and a half in length, when the active formation of new substance takes place, and when the trees get ready, by the storage of reserve materials, for the period of the winter's rest. Even on August 30, we noted the display of red autumn leaves on the red-maple, Acer rubrum, and on the sour-gum tree, Nyssa sylvatica. A few cases of special interest should be mentioned before considering the table in detail. The pitch-pine, Pinus rigida, begins to show signs of staminate flower development on May 5. On May 20, these flowers discharged pollen. Presumably, the pollination took place on that date, or before the end of May. The pollen-grain, or microspore, is drawn into the pollen-chamber, where it remains over the summer, autumn and winter, until the next spring. The pollen-tube is probably sent out into the nucellus, as soon as the grain is deposited, but it stops active growth until April of the next year. During April the pollen- tube begins to renew its penetration of the nucellus, and the large tube- nucleus enters the pollen-tube, where it is invested completely by starch grains, and at the same time the generative cell divides into stalk-cell and body-cell. The pollen-tube consumes about two months in travers- ing the nucellus after its second start, entering the archegonium about July I. Just before fertilization the body-cell divides to form the two male cells, so that at this time the pollen-tube contains the tube-nucleus, stalk-cell nucleus and the two male cells. The tip of the pollen-tube, having penetrated the overlying cells of the nucellus, reaches the wall of the embryo-sac, and either passes directly through it, or flattens out upon it in a foot-like expansion, sending out a small branch to reach the neck of the archegonium. The neck-cells are crushed and the pollen- tube comes in contact with the egg about July 1, when fertilization occurs. While the seeds are developing the cones rapidly enlarge until they reach mature size by September 6, for cones brought into a warm, dry laboratory that were gathered on that date opened and discharged their seeds after being kept indoors for a week. Outside, however, the cones do not open until the warm days of Indian summer, for a trip to the pine-barrens on November 13, 1912, showed many, but not all, of the cones of the pitch-pine fully open and discharging their winged seeds. Some observations of Brown * are apropos. He finds that in the pinery of Cornell University the growth of the cambial layer in young twenty- to thirty-year-old specimens of Pinus rigida began in the vicinity of * Brown, Harry P.: Growth Studies in Forest Trees — (i) Pinus rigida Mill. The Bo- tanical Gazette, Liv: 386-401, November, 1912. '4 2 10 VEGETATION OF THE NEW JERSEY PINE-BARRENS Ithaca as early as April 15, and it is probable also that in the vicinity of Ithaca growth begins at about the same time each spring. As the growing season is shorter at Ithaca than in the pine-barrens, it is prob- able that growth of the cambial layer of the pitch-pine begins early in central New Jersey. Growth at Ithaca began first in the twenty- to thirty-year-old specimens at some distance below the apical shoot, but during a period of nineteen days gradually spread upward until it reached the apex of the trees. Growth spreads down the main axis faster than it does along the lateral shoots. Except in the terminal shoot, growth in diameter was more rapid between May 25 and June 6. In the terminal shoot itself greatest rapidity of growth was manifested between June 6 and June 1 5. Brown finds also that there is no appreci- able difference in the time of cambial awakening on the north and south side of the trees. The evergreen flowering-moss, or pyxie, Pyxidanthera barbulata, also claims attention. The earliest date which I have of open flowers is March 23, and the last date, when only a few flowers were found, is April 30, but the herbarium records given by Bayard Long extend the flower period of this plant to May 10, so that the total period of flowering of this species is forty-eight days. On June 13 fully ripe capsules and seeds of this species were found, so that eighty-two days elapsed from the opening of the first flower until the development of mature fruit and seeds. The period of flower and fruit development in some pine-barren plants is much shorter than this, for example, Comptonia asplenifolia, eighty days, Lupinus perennis, seventy days, Acer carolinianum, sixty- five days, Hudsonia ericoides, sixty days, Salix humilis, forty-seven days. Longer periods than eighty-two days are found in quite a few plants of the pine-barrens. The white water-lily, Castalia odorata, flowers from May 23, the earliest date that I have on record, to October — about one hundred and thirty days. The production of fruit begins in this plant early in July and continues into October. The seeds may germinate immediately after escaping from the fruit, or may remain dormant in cold water until the following spring: the first is probably the case with fruits ripened before September, the second with those ripened later, according to Conard. Conard* found great numbers of seedlings (var. minor) in a pond south of Marlton, N. J., on August 30, 1900, with only submerged leaves. The first leaf is filiform, about 1 .6 centimeters long. The second is elliptic-oblong. Intermediate leaves are ovate, cordate, rounded at apex, with rounded basal lobes and sinus. Late submerged * CoNARU, Henry S.: The Waterlilies, 1903: 181. VEGETATION OF THE NEW JERSEY PINE-BARRENS 211 leaves are sagittate-cordate, with deep, narrow sinus, are stiff in texture, bright emerald,, and stand erect or semi-erect. The actual period of flowering is short for the amentiferous trees and shrubs, as is shown by giving the data for the following trees, arranged systematically: Populus grandidentata 12-13 days Salix nigra 20 " humilis '5 " tristis 15 Myrica carolinensis 35 Comptonia asplenifolia 20 Hicoria glabra 20 Betula populifolia 25 Alnus rugosa 25 Quercus coccinea '5 velutina 17 triloba 16 |' " ilicifolia 17 " marylandica 17 phellos 20 alba 20 stellata 15 '' prinus 15 " prinoides 18 But in the majority of these trees and shrubs, with the exception of Populus grandidentata, which matures its fruit in May or June, and of Salix nigra, S. humilis, and S. tristis, which mature their fruits by the middle of May, all other trees and shrubs show a slow development of the fruit, which is not mature until some time has elapsed. The fol- lowing facts are known about the maturation of the fruit of the remain- ing species in the list given above. The fruit of Myrica carolinensis matures between the end of July and the middle of August. That of Comptonia asplenifolia matures between the middle of June and early July. The fruit of the pignut, Hicoria glabra, white-birch, Betula popu- lifolia, alder, Alnus rugosa, do not mature unti late August and Sep- tember. The oaks of the pine-barrens may be divided into two groups. The annual-fruited, or white-oaks, comprising Quercus alba, Q. stellata, Q. prinus, and Q. prinoides, mature their fruits in September of the first year, i. e., the year of pollination and fertilization. The biennial- fruited, or black-oaks, comprising Quercus coccinea, Q. velutina, Q. falcata, Q. marylandica, Q. phellos, and Q. ilicifolia, do not mature their fruits until September of the next year after pollination. For the small herbaceous plants the period of flower production of the several pine-barren species of Polygala is a long-drawn-out one. With the exception of 12 species, all of the 42 species of the family Com- 212 VEGETATION OF THE NEW JERSEY PINE-BARRENS positae, represented in the pine-barren flora, flower after the middle of July, and most of them in August and September. We will now tabu- late the number of plants that come into flower at intervals of five days throughout the year, so as to ascertain at what season the majority of pine-barren plants come into flower. We will then be able to decide whether the vegetation, as far as flowering is concerned, is a spring, summer, or autumn vegetation. TABLE SHOWING NUMBER OF PLANTS THAT COME INTO FLOWER AT FIVE-DAY INTERVALS FROM MARCH 15 TO SEPTEMBER 10, INCLUSIVE March I o o o 2 2 2 ^ . . . lO I c 6 ] 20 . . 2S April I 3 c 5 o lO . I c 33 ] 2 20 23 2=; . ^ 0 May I . . . . . f 0 i: 3' 4 10 IS 91 i 21 20 32 3 2"; . . . . . June I f 3 c. 30 17 10 I c H3 30 29 I 4 20 . 25 July 5- 10. 15- 20. 25- August I . 5- 10. •5- 20. 25- September i . 5- 10. 15- 20. 25- October I . 5- 10. 15- 20. 25- 106 I 30 24 29 72 24 9 24 15 o 15 I 10 3 o 1 Schizaea o It will be seen from the table that the blossoming of pine-barren plants is irregular, and a graphic curve of the phenomena illustrates the same thing. The period extends from March 15 to September 10, approxi- mately six months. If, however, we take the number of plants that bloom by monthly periods, we find that the majority of pine-barren plants flower during May, June, July, and August, with the largest number during June, July, and August. In these four months 382 species out of 436 species come into flower. But to reach a true statement of the number of plants in flower during any one month we must refer again to our graphic tabulation of the pine-barren species and count the VEGETATION OF THE NEW JERSEY PINE-BARRENS 213 number actually in bloom during the successive months. The follow- ing is the result of this enumeration: Number Number OF Plants that Came Month in Flower into Bloom March 6 6 April 41 33 May 128 91 June 198 113 }^y 247 106 August 263 72 September 207 1 5 October 58 o A comparison of the figures in the two columns of the preceding table indicates that although a certain number of plants may have come into flower in a certain month, yet many of these plants continue the period of flowering into the next following month, or even to two or three months, so that the number of plants actually in bloom in any one month is augmented considerably, as shown in column one. For example, many of the early spring flowers go out of flower early in June and many late summer blossoming plants begin to flower in June, so that both early spring and late summer plants help to make the total June number. This applies also to July, August and September. As no plants come into flower after September lo and only 15 before that date, the large number of plants in flower in September is due to the continuation of the flowering periods of plants that blossomed earlier. The same applies particularly to October, a month in which no plants blossomed, but in which we fmd 58 species in actual flower. However, the important point which the foregoing table demonstrates is that the months of June, July, August and September are the flower months of the pine- barren vegetation season. The month of May ought to be included, perhaps, so that practically five months may be looked upon as the ac- tive flowering season of pine-barren vegetation with the culmination of the flower display in August. From March 15 to October 30 the pine-barren region is practically never without some plant in flower. March, April, and October are the dullest months, and November, December, January, and February com- prise the dormant period. Before concluding this account of the phenol- ogy of pine-barren vegetation we will construct a few figures to illustrate the procession of vegetation in each of the principal plant formations of the region, viz., pine-barrens, cedar swamps, deciduous swamps, ponds, plains, and savannas (Fig. 1 56). The procession of flower development as illustrated by a few plants of these formations is as follows: 214 VEGETATION OF THE NEW JERSEY PINE-BARRENS Figure 156 Procession of flower development in a series of plants of the pine-barren, cedar swamp, deciduous swamp, pond and plains formations. VEGETATION OF THE NEW JERSEY PINE-BARRENS 215 Pine-Barren Formations: Epigaea repens April 5-May 10. Helianthemum canadense May 10-July 25. Andropogon scoparius •July 20-October 10. Bog Formation: Orontium aquaticum March 25-June 5. Pogonia ophioglossoides June 5 — July 10. Hypericum boreale July la-September.io. Gentiana porphyrio September lo-October 15. Deciduous Swamp Formation: Acer carolinianum March 2(>-April 20. Vaccinium corymbosum April 20-May 25. Magnolia glauca May 20-July 15. Polygala brevifolia July 15-October 15. Pond Formation: Orontium aquaticum March 2 5-June i . Castalia odorata June i-October 1. Plain Formation: Pyxidanthera barbulata March 23-May 10. Andromeda mariana May 5-July 5- Pteris aquilina July 5-September 20. Savanna Formation: Alnus rugosa March 1 5-April 10. Chamaecvparis thyoides April i-April 20. Pvrus arbutifolia April 20-May 25. Iris prismatica May 20-June 25. Lophiola aurea June 25-July 25. Eriocaulon decangulare July 15-October 5. It will be noted that two plants cover the flowering period of pond vegetation; three the period for the pine-barrens and plains respectively; four the white-cedar swamp and marsh periods, and five the period of flowering in the savannas. Graphically the procession in each of the formations may be indicated by a circle in which the size of each seg- ment of the circle represents the relative length of time at which the given species is in flower (Fig. 1 56). CHAPTER XVI VEGETATIVE PROPAGATION AND STRUCTURE OF THE SHOOTS . Much attention has been given of late years to a study of those organs of the plants which are concerned with the absorption of food, which serve to anchor the plant, which store food, or which renew the plant by vegetative propagation. As the pine-barren vegetation is an ancient and exclusive one in the region where it is found and exists in climax form, it is important to study the organs by which these plants have been able, in undisturbed condition, to withstand the encroachment of other nearby formations which have reached also a climax condition. The pine-barren species occupy the soil layers so closely with their over- ground and their underground parts that few new plant introductions have succeeded in making their way into such closed formations which have occupied the region since post-Miocene times. Abroad several plant ecologists have emphasized the importance of this examination of the propagative shoots and roots. Only a few of the papers will be mentioned in passing. Tansley and Fritsch,* in a study (1905) of the flora of the Ceylon littoral, give some attention to this subject. Warm- ing,! in 1908, published his careful investigations on the structure and biology of arctic flowering plants, and in these studies he was assisted by Petersen. Yapp,J in the same year, gives an excellent synopsis of the underground parts of the plants of Wicken Fen, and in a later, more- detailed paper, the results of his research on the stratification in the vegetation of a marsh and its relations to evaporation and tempera- ture. Ostenfeld, in the Botany of the Faroes (1908: 867-1018), con- siders the land vegetation of these islands with special reference to the higher plants and the vegetative propagation and structure of the shoots. Eugen Hess,§ in an inaugural dissertation, discusses the special structure * Tansley, A. G., and Fritsch, F. E.: The Flora of the Ceylon Littoral. The New Fhytologist, iv: 1-17; 25-55. t Warming, Eugen : The Structure and Biology of Arctic Flowering Plants. Meddelel- ser om Gronland, xxxvi. t Yapp, R. H.: Wicken Fen. The New Fhytologist, vii: 61-81; On Stratification in the Vegetation of a Marsh, and its Relations to Evaporation and Temperature. Annals of Botany, xxiii: 275-319, April, 1909. § Hess, Eugen: Ueber die Wuchsformen der alpinen Gerollpflanzen, Universitat Zurich, 1909. 216 VEGETATION OF THE NEW JERSEY PINE-BARRENS 217 of the alpine boulder plants. Sherff* has pursued much the same line of investigation in his studies of the vegetation of Skokie Marsh, with special reference to subterranean organs and interrelationship. More recently Yappt has published a paper on a single plant, Spiraea ulmaria, and its bearing on the problem of xeromorphy in marsh plants. Can- non,! in two publications of considerable length from the Desert Bo- tanical Laboratory at Tucson, has described the root habits of desert plants. Late in my studies of pine-barren vegetation 1 determined to under- take a study of the underground parts of a series of pine-barren species. The result of that study is given in the following pages for 32 pine-barren plants arranged systematically. Some few of the descriptive details and some of those shown in the drawings were obtained from herbarium material which served also to check the observations made upon fresh parts while in the field. The hapaxanthic species, or those that only flower once, propagate by seeds alone, whereas many of the perennials propagate both by seeds and vegetatively. Ostenfeld has dealt with them under three categories: 1. Spot-bound (sedentary) species, i. e., species which have no stolons, creeping rhizomes, nor bud-producing roots, or the rhizomes are so short that the plants have little, if any, power of wandering vegetatively. 2. Wandering species with epiterranean (above ground) runners. 3. Wandering species with subterranean shoots, stolons, creeping rhizomes, or bud-producing roots. Hess,§ in the important paper mentioned above, has classified the different kinds of shoots. As his classification is applicable to the study of pine-barren plants, a translation is given by way of an introduction to what follows. To bring out the salient features of this system pine- barren species have been arranged as follows: 1. Spot-Bound Shoots A. Single Spot-bound Shoots 1. We may designate simple, upright, terrestrial stems that bear a crown of leaves at the top, namely, caudices. The primary root persists, * Sherff, Earl E.: The Vegetation of Skokie Marsh, with Special Reference to Sub- terranean Organs and their Interrelationship. The Botanical Gazette, Liu: 415-435. t Yapp, R. H.: Spiraea ulmaria and its Bearing on the Problem of Xeromorphy in Marsh Plants. Annals of Botany, xxvi: 815-870. J Cannon, W. A.: The Root Habits of Desert Plants. Publ. No. 131, Carnegie Insti- tution of Washington, 191 1; Some Features of the Root-Systems of the Desert Plants. Popular Science Monthly, July, 1912. § Hess, Eugen: Ueber die Wuchsformen der alpinen Gerollpflanzen, pages 38-49. 2l8 VEGETATION OF THE NEW JERSEY PINE-BARRENS but secondary adventitious roots may be formed. Pinus rigida, Quercus marylandica, Cassia chamaecrista, Chimaphila umbellata, Melampy- rum americanum. 2. Under this head may be included bulbous shoots. Cypripedium acaule, Habenaria blephariglottis, Eupatorium verbenaefolium, Liatris graminifolia, L. pilosa, Hieracium venosum. B. Spot-bound Shoots in Clusters 3. By the branching of spot-bound shoots are formed aerial tufts or cushions. Few species form only tufted shoots without adventitious roots. Frequently, however, adventitious roots are not formed as an occasional adaptation of such species which normally grow with other plants. The rounded cushions in thick crowding display radially sepa- rated branches. Quercus prinoides, Baptisia tinctoria, Rhus copallina, Hudsonia ericoides. 4. The tufts of branches, which are provided with roots, abundant at times, are illustrated by the grasses and sedges. Such shoots with only sufficient space to grow upright develop from a short, horizontal piece. Uninterrupted growth leads to the covering of extensive surface, so that cushions are formed, but such cushions are formed of parallel shoots, not radial. 5. Caudex multiceps. — Uniform extended stems of mono- or sympodial branching, which grow over or under ground, were termed by Areschoug terrestrial stems (Erdstamme). If the branch grows vertically, or sharply upright, it reaches the surface in a single tuft of leaves. Hitch- cock designates such a simple vertical shoot a caudex. A much- branched, upright terrestrial stem bearing tufts of leaves at each ex- tremity is designated by Hess a branched caudex {Caudex multiceps). Hess distinguishes between the spot-bound form and the spread-out form, which Hitchcock calls crown (Krone), and Warming a multicipital root (radix multiceps). Whether these terrestrial stems have primary or adventitious roots or whether the flower-stem is leafy or not, is, there- fore, indifferent. Examples are Euphorbia ipecacuanhae, Kalmia lati- folia, Tephrosia virginiana. 11. Spreading Shoots Spreading shoots are either normal shoots or else they are elongated runners. A. Spreading Shoots of Normal Form 6. Hess calls aerial shoots of this group which develop without roots crested shoots (Schopftriebe). Ratzeburg calls them periwigs (Peruck- VEGETATION OF THE NEW JERSEY PINE-BARRENS 219 en), or wig forms. In gravel, the branches are not aerial, but grow partly in the light, partly in the dark, of the soil spaces, and the small leaves on them are generally colorless. Sometimes the wig shoots wind about, between, and over the stones, a distance of 20 to 30 centi- meters— Pyxidanthera barbulata. 7. Shoots which spread radially as branches of a single stem, and which thus form a crown, may be termed wheel shoots, or simply radiate shoots. These radial shoots do not develop roots except unusually long, hence they should never be classed as rhizomes. Hudsonia tomen- tosa, Arenaria caroliniana. Cassia chamaecrista. 8. The surface-spreading shoots which develop adventitious roots may be termed turf-forming shoots. Such are termed generally by American botanists creeping stems, and they may be orthotropic or plagiotropic. 9. Rhizome (rootstock) is any horizontal or oblique perennial stem which grows on the ground, or in it, and produces roots and shoots that are usually leafy and green. Pteris aquilina, Sparganium eurycarpum, Smilax glauca, Comptonia asplenifolia, Myrica carolinensis, Hypericum densiflorum, Nymphaea advena. Ilex glabra, Asclepias obtusifolia, Gaultheria procumbens, Andromeda mariana, Vaccinium vacillans, Arctostaphylos uva-ursi, and Sericocarpus asteroides. B. Extension by Elongated Slender Shoots 10. Wandering shoots (Wandertriebe) are similar to the turf -forming shoots, but they are hidden beneath the surface and grow through the interstices of the earth, or between the stones, if the soil is a gravelly or a stony one. When they reach the surface and grow upon it, they can- not be distinguished from the turf-forming shoots. Tofieldia racemosa, Pyxidanthera barbulata. 1 1. The slender running shoots (Laufen) which strike root at the tip are called stolons, or runners. An offset is a short stolon. 12. A sucker is an ascending stem arising from a subterranean base usually connected with the mother plant. It is not included in the classification which Hess has given, because it does not occur among the plants which he has studied, viz., the rubble, or boulder plants (Geroll- pflanzen). Detailed Root and Stem Studies It is only within recent years that plant ecologists have directed their attention to the study of the root and the underground stem portions of plants, as well as their physiologic condition. But the prospect is fair 220 VEGETATION OF THE NEW JERSEY PINE-BARRENS that in a few years we will have definite information on this important subject. The roots and underground stems of pine-barren plants are of interest because of their relation to the perennation of the species and to the physiologic activities of the shoots. There is a close relation between the character of these parts and the distribution of the plants, the association of the different species in the different plant formations, the formation of the leaves, of the flowers, and the commencement of new growth. According to Cannon, the work of Rimbach, Biisgen and Freidenfeldt, as reviewed by von Alter (Wurzel-studien, Bot. Zeit., 67: 175, 1909), is important because their researches indicate that the root system of flowering plants may be separated into two groups, according to the character of the terminal roots: they are either intensive or ex- tensive. Intensive root systems have fine terminal roots; they are branched richly and occupy relatively small soil volume. Extensive root systems have coarse ultimate rootlets, are not richly branched and occupy a relatively large soil volume. Cannon * describes three main types of root systems to be found in the desert plants of the southwestern United States: (i) Root systems which extend horizontally from the main axis and lie, for their whole course, near the surface of the ground. (2) Root systems which are characterized by a strongly developed tap- root going down directly to a depth determined in part by the character of the soil, in part by the penetration of the rains, and in part by the character of the root itself. (3) Roots that not only reach widely, but also penetrate fairly deeply. Where the root system is an obligate type the distribution of the species is much restricted, but where it undergoes modification with changed environment the distribution of the species is much less confined. It is of interest to note especially that, as a rule, it is the latter kind of root system that is developed by such plants as occur where the soil conditions are most arid, from which it follows that the generalized type of root system (No. 3 above) is really the xerophytic type par excellence and not the type with the most deeply penetrating tap-root, as might be supposed. Reference will be made to these points as we proceed with our studies. Having classified the pine-barren plants as to their underground parts, it is important to describe next in detail the stem and root systems of each of these plants. Pteris aquilina (Bracken-fern). — This fern develops a brown under- ground stem, or rhizome, which in the specimen studied reached a length of 6.7 meters (22 feet). The usual thickness is about 1.27 centimeters * Cannon, W. A.: Some Features of the Root Systems of the Desert Plants. Pop. Sci. Month., July, 1912; Botanical Features of the Algerian Sahara. Publ. 178, Carnegie Institution of Washington, 1913. VEGETATION OF THE NEW JERSEY PINE-BARRENS 221 {^A inch), and along each side run Hghter bands called the lateral lines, which probably function in the respiration of the rootstock. The grow- ing-point, which is rounded, whitish, and covered with paleae, has a single apical cell, and the stem branches in two ways — first, by apparent dichotomy of the terminal growing-point, and second, by the formation of adventitious buds. The latter are produced singly on the dorsal side of the leaf-stalks near the base. The bases of the old leaf-stalks persist for a few years, but they ultimately disappear. The roots, like the stem, develop by means of an apical cell which is covered with a root-cap. Root-hairs are found. As to the structure of a single rhizome 6.7 meters (22 feet) long, the following notes are given in the metric units and the English units of measurement. No. OF Branch 1 . 2. 3 4- 5- 6. 7- 8. 9 10. 1 1 . 12 •3 14 •5 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 Distance behind Apex Metric 5 23 4.6 6.8 9.1 1-4 1.6 1.9 2.0 2-3 2-5 2.7 2.9 3-2 3-3 3-3 3-7 3-9 4,0 4.2 4-3 4-5 4-7 4.8 5.0 5-2 5-4 5.6 5-7 5.8 6.0 6.0 6.2 6.4 cm. dm. English I ft. 3 " 3 " 4 " 5 " 6 " 6 " 7 " '3 >3 14 14 ■5 15 16 17 17 18 19 19 19 20 20 21 9 6 " 3 " o " 9 " 6 " 3 10 6 " 2 10 " 6 " 6 " o " 6 " 4 10 " 4 10 4 10 4 10 " 5 1 " 8K" 8>^" I in. 6^4. ins. 9 " o " 6 " 10 " Length of Branch Metric 2.5 cm. 6.3 " i.o dm. 17.0 dm. 12.0 dm. 3 dm. 3 cm. o 5 5 5 5 o dm. 7 " 3 cm. 3 " English I in. jyi ins. 4 ins. 7 ins. 5 ins. I ft. 9 ins. 2}/^ ins. 2 " 3 " 3 " 3 " 3 " I ft. 7 ins. 2>^ " 2>^ " rhizome rotten 222 VEGETATION OF THE NEW JERSEY PINE-BARRENS The underground rhizome of the bracken-fern may be classified with the extensive kind of von Alten, and to the first division of Cannon's clas- sification, and the plant is a hemicryptophyte, according to Raunkiaer. Figure 157 Plotted rhizome of Pteris aquilina. I small square = I inch. A, Level of soil. B, Rhizome (true depth), upper dotted lines. branches (lower dotted lines). C, Branches. D, Dead The depth at which the rhizome was found varied, as it accommodated itself in its growth to the larger pine and oak roots under which, or over which, it grew in its course through the soil. At 22.8 cm. from the apex it was 1 1.4 cm. deep. " 30.4 " " " " " " 13.9 " 60.8 " " " " " " 12.7 " " 152.0 " " " " " " 8.8 " " 182.4 " " " " " " 20.3 " " 243.2 " " " " " " 20.3 " " 295.4 " " " " " " 8.8 " " 362.3 " " " " " " 17.7 " " 443.3 " " " " " " lo.i " " 547.2 " " " " " " 11.4 " " 577-6 " " " " " " 10. 1 " " 608.0 " " " " " " 15.2 " The conspicuous parts of this fern are large green fronds which arise above the ground to a height of 45 centimeters (18 inches) to 60 centi- meters (2 feet), occasionally 15 to 18 decimeters (5-6 feet), and consist of a rachis with thrice pinnate divisions. The rachis of the frond may be followed some distance into the ground. Its buried portion is brown in color, and it is attached to the rhizome below, from which true roots are given off. Traced in one direction from the attachment of the frond the rhizome exhibits the withered bases of the fronds, developed in former years, which have died down; while in the opposite direction it ends, sooner or later, in a rounded extremity covered with fine brown hairs, which is the growing-point of the stem. Between the free end and the fully formed frond rounded projections, the rudi- ments of fronds, are seen. These will attain their full development in the following years (Fig. 157). VEGETATION OF THE NEW JERSEY PINE-BARRENS 223 Pinus rigida (Pitch-pine). — This pine is a typic phanerophyte, ac- cording to the classification of Raunkiaer (Figs. 158, 159, 160). It develops a powerful tap-root, which persists for many years unless, as in the plains, it comes in contact with a hard, impervious subsoil, when it rots away. Sometimes it is bifurcated, as in a tree studied along Shark River Bay. Strong lateral roots are developed from the main primary root. The strongest of these sec- ondary roots are near the soil surface, in a specimen measured 10 centimeters below the surface, and they run horizontally for some distance, finally sinking obliquely downward, or at times sharply at right angles into the deeper soil levels. Several of these powerful lateral roots may be formed at slightly different depths. The smaller and shorter secondary roots are developed from the lower part of the primary root, and from a physiologic standpoint may be considered to be a part of it, while the more superficial and stronger secondary roots belong in func- tion to a slightly different category. To give a few specific examples: A tree was studied at Como Lake. Its first large su- perficial secondary root was given off 10 centimeters below the soil surface, where it soon bifurcated. The lower division sank to a level of 23 centimeters and ran a dis- tance of 4.5 meters; then it grew obliquely downward to a depth of 1.5 meters, reach- ing a distance of 6.5 meters measured at the soil surface from the base of the tree. Another horizontal root grew down to a depth of 1.7 meters below the soil surface. Some of the shorter lateral roots arose 30 centimeters below the surface of the soil. In a low pine tree studied in the Upper Plain, which was only 30 centimeters tall, it was found that the largest roots were horizontal and superficial, gradually dipping to a gravel layer 50 centimeters below the surface. The primary root had decayed 30 centimeters below the surface. This decay had been hastened by the root growing to the hard pan, which has been described previously. Figure 158 Pinus rigida, showing root system. 224 VEGETATION OF THE NEW JERSEY PINE-BARRENS Sparganium amerkanum (Slender Bur-reed). — This marsh plant is found in the pine-barrens near Forked River, Bamber, Tuckerton, Park- dale, Bear Swamp and Clementon. It is a cryptophyte, according to Raunkiaer's classification, being included in the division of helophytes along with Typha and other marsh plants. The rhizomes of this plant are formed about two centimeters below the surface, and are strong plagiotropic shoots with large terminal buds. According to Sherff, who has studied S. eurycarpum, such plants with strong, thick rhizomes have a decided advantage over other plants of less vegetative vigor. Dulichium spathaceum. — If we examine the underground part of Dulichium, we find a rather long horizontal rhizome with sympodial ramifications. The internodes are distinct and covered partly with rudimentary sheathing leaves. Relatively strong roots develop above the nodes, especially from the lower surface. Several stems arise from Figure 159 Figure 160 Horizontal plan of the root system of the Horizontal view of root system of pitch- pitch-pine, Pinus rigida. o, Sinking pine pine, roots; -, horizontal pine roots, as shown at tips of branches. the rhizome, and they show a structure very different from what we are accustomed to see in the Cyperaceae, having numerous cylindric and hollow internodes. The leaves of Dulichium are either reduced tubular sheaths upon the rhizome and at the base of the stems, or they are de- veloped normally with a linear blade, a short, crescent-shaped ligule, and a long tubular sheath. The plant is a hemicryptophyte. Andropogon scoparius Michx. (Beard-grass). — This grass, which has tufted culms 4 to 12 decimeters tall, is a typic hemicryptophyte. The leaves and culms arise from a rounded, woody base and the filiform sec- ondary roots spread in every direction (Fig. 161). Panicum depauperatiim. — This erect or ascending grass, 2 to 4 deci- meters tall, occurs in the driest situations in the pine-barrens. Its tufted culms and leaves arise from a plate-like stem provided with nu- merous filiform secondary roots. A hemicryptophyte (Fig. 162). VEGETATION OF THE NEW JERSEY PINE-BARRENS 225 Ahama (Narihecium) americaniim. — The bog-asphodel has an aerial shoot, 2.5 to 4 decimeters tall, which arises from a horizontal rhizome covered with the fibrous remains of the old leaf-bases. The secondary roots are numerous, arising from the under surface of the rhizome. It is abundant in the wet savannas along the Wading River. A hemicryp- tophyte (Fig. 163). Xerophyllum asphodeloides. — The turkey's-beard is a characteristic plant of the New Jersey pine- FlGURE 161 Andropogon scoparius. Figure 163 Abama americana. Figure 162 Panicum depauperatum. barrens. Its long, needle-like leaves form a dense hassock which repre- sents the massing together of the leaves of several distinct plants that have a short rhizome beset with numerous secondary roots. The leaves are flattened at the base and overlapping. A hemicryptophyte (Fig. 1 64) . Tofieldia racemosa. — The false asphodel is a slender perennial, lilia- ceous plant, mostly tufted, with a slender plagiotropic rhizome growing about I centimeter below the surface of the boggy places in which it occurs. The rhizome shows tufts of roots at the nodes, reaching up nearly to the surface, and a few superficial, inordinately long wandering 226 VEGETATION OF THE NEW JERSEY PINE-BARRENS roots which develop a tuft of smaller roots at their extremity. The flowering scape of this plant reaches a height of 3 to 9 decimeters. A hemicryptophyte (Fig. 165). Smilax glauca. — The saw-brier is an erect or climbing plant that pro- duces a tough, wiry, plagiotropic rhizome, white in color, divided into clearly marked nodes and internodes. The nodes bear pointed buds protected partially by the sheathing leaf-base, which is the only part of the leaf found below ground. Two kinds of roots are formed, namely, long nodal roots that may reach a length of 17.7 centimeters (7 inches), and short internodal roots. A hemicryptophyte (Fig. 166). Cypripedium acaule (Moccasin Flower). — This beautiful orchid has a short, rounded stem just below the surface of the soil. From it arises a cluster of slender roots from 3 to 25 centimeters long, produced in large numbers and of a brown color. Two large, oval leaves surround the Figure 164 Xerophyllum asphodeloides. Figure 165 Root system of Tofieldia racemosa. Figure 166 Smilax glauca. base of the single-flowered scape. The fibrovascular bundles of old leaves in a shredded, fibrous condition are seen frequently at the base of the stem and on the surface of the soil. A hemicryptophyte (Fig. 167). Pogonia ophioglossoides* — This is a terrestrial orchid with a slender rhizome, which is relatively short, vertical, or ascending, and densely covered with long, unicellular hairs, a unique feature on underground stems. The rhizome in this species, according to Holm, consists of five * Cf. Holm, Theo.: Pogonia ophioglossoides Nutt., a Morpiioiogical and Anatomical Study. American Journal of Science, IX : 13-19, January, 1900. VEGETATION OF THE NEW JERSEY PINE-BARRENS 227 Figure 167 Cypripedium acaule. distinct internodes, the uppermost passing into a flower-bearing stem. The rhizome is vertical, bearing rudiments of two green leaves, and two scale-like and membranaceous, while a third scale-like leaf is still fresh and surrounds the base of the flower-bearing stem. The monopodial growth is characteristic of this rhizome, which ceases, however, when the flower- bearing stem dies away. After that time it be- comes a sympodium, as the axillary bud will de- velop and continue the growth of the rhizome. This is, however, not the only bud which is visible, since another, though dormant, is developed in the axil of another leaf. This bud will grow out into a branch if the upper part of the rhizome becomes injured, but not otherwise. The main propaga- tion of this orchid is by means of root-shoots. This striking feature makes a study of the roots of great interest. It appears that the shoot de- velops terminally at the end of the root, and a secondary root pushes out from the base of the young shoot and in the same direction as the mother root, as if it were a continuation of it. This secondary root repeats the development of another shoot in exactly the same manner as the first. The other secondary roots which develop higher up on the shoot grow often from the beginning in a down- ward direction, and do not form new shoots, but are concerned probably with absorption. Hyphae are found in the hypoderm and in the cortex of these roots. A hemicryptophyte. Limodorum tuberosum {Calopogonpulchellus) is an orchid which has a 4- to 12-flowered scape arising from a solid bulb, which is sheathed below by the base of the solitary grass-like leaf. The cortex of the slender roots of this orchid is supplied with mycorhizal hyphae, suggesting that it and Pogonia are not entirely autophytic, but hemisaprophytic. Habenaria blephari glottis (White Fringed Or- chid).— This orchid, found in white-cedar swamps, has two kinds of roots arising from the base of the stem, namely, slender feeding roots and elongated tuberous roots, from which the buds that develop into new shoots arise. The stem which arises from these clustered roots is 4 to 6 decimeters tall, and bears Figure 168 Underground system of Habenaria blephariglot- tis. 228 VEGETATION OF THE NEW JERSEY PINE-BARRENS several linear, lanceolate leaves on its lower part, and small linear leaves just below the crowded raceme of pure white flowers. A hemicrypto- phyte (Fig. i68). Comptonia asplenifolia (Sweet-fern). — This low shrub, 3 to 6 deci- meters tall, with sweet-scented, fern-like foliage, belongs to the group of nanophanerophytes of Raunkiaer. It arises from a long horizontal, Figure 169 Comptonia asplenifolia. Figure 170 Horizontal view of the root system of Quercus marylandica. Figure 171 Quercus marylandica. frequently forking root, sometimes three feet long, and about the thick- ness of a lead-pencil. It produces filiform lateral roots and is found in the surface soil just beneath the layer of pineneedles (Fig. 169). Myrica carolinensis. — The bayberry is a nanophanerophyte. It de- velops ordinarily a vertically descending tap-root, but where the leaf mold, consisting of pine needles and other leaves, has been piled in masses, the bayberry develops a horizontal rhizome the thickness of a VEGETATION OF THE NEW JERSEY PINE-BARRENS 229 lead-pencil. It runs a distance of about 75 centimeters away from the main stem, and sends down short lateral roots, which become associated Figure 172 Ouercus ilicifolia. intimately with the leaf mold, so that it is almost impossible to separate them. 230 VEGETATION OF THE NEW JERSEY PINE-BARRENS Quercus prinoides (Dwarf Chestnut-oak). — This low, bushy oak de- velops a strong tap-root, 3 to 4 centimeters thick, that grows deep into the soil. The stronger lateral roots are found some distance below the surface. In one specimen studied the first strong lateral was 30 centi- meters below the surface, the other, 42.5 centimeters. The smaller lateral roots arise from the primary root just below the soil surface. In a tree studied in undisturbed pine-barrens four stool shoots were found growing at first prostrate beneath the soil and then rising at the ends 60 centimeters above it. A nanophanerophyte. Quercus marylandica (Figs. 170 and 171). — The black-jack oak grows to be a small tree in the pine forests of New Jersey. In a specimen 12 decimeters tall was found a conic tap-root descending vertically into the soil. From this central root developed two kinds of lateral roots. First, short roots are borne on the primary root at various depths below the surface. Second, long and thick lateral roots arise at different levels. In the tree studied there was one 9 decimeters long, 18 centimeters below the surface; a second, 5.5 decimeters long, was found 25 centimeters below the surface; a third grew 4 decimeters below the surface. These are formed usually in different ranks, so that they do not stand one above the other. Quercus ilicifolia. — The bear-oak is a dwarf tree i to 3, rarely 6, meters high, usually covered with acorns in the fall of the second year. The stems arise from a gnarled, knotted, and twisted crown, and the strong roots are several in num- ber, descending obliquely into the upper subsoil. Horizontal roots are found in the upper soil layer near the surface. A mesophanerophyte (Fig. 172). Sassafras variifolium. — The sassafras is usually a small tree in New Jersey which may reach a large size elsewhere (a phanerophyte). In the young trees in the pine forests of New Jersey the stems one meter tall arise from a horizontal rhizome sparingly provided with roots (Fig. 173). Comandra umhellata. — The bastard-toadflax is a hemicryptophyte. It has a slender horizontal rhizome and filiform descending roots. The plant, which has upright stems 15 to 30 centimeters tall, is probably parasitic on the roots of other plants. Hudsonia tomentosa. — This heath-like, low plant has a chocolate- brown tap-root growing downward vertically a length of 12.5 to 1 5 centi- meters. The lateral roots which are developed from the point where the Figure 173 Sassafras variifolium. VEGETATION OF THE NEW JERSEY PINE-BARRENS 231 Figure 174 Hudsonia tomentosa. lateral branches arise are 10 to 15 centimeters long and wiry. The lateral branches which radiate outward from this central point are at first plagiotropic and at the tips orthotropic. Hence arises a cushion- like plant of a grayish-green color covered with scale-like, alternate, persistent, downy leaves. Hence we have a plant which has shoots arranged in a wheel-like or radiate manner (Fig. 174). Hudsonia ericotdes.— This perennial plant, with awl-shaped leaves, arises from a superficial rhizome about the thickness of a lead-pencil. Several strong roots and a lot of filiform ones are developed. Several stems may grow from the same rhizome. A hemicryptophyte (Fig. 175). Hypericum densiflorum.— This is a bushy shrub, 0.5 to 2 meters tall, and branches richly above the ground, with slender branches crowded with small leaves. The root is arched or horizontal, with numerous adventitious roots growing down from the under surface of the horizontal portion. The main root, which is a direct continuation of the horizontal portion, grows downward obliquely, where it divides into a number of strong branches. A short stub on the upper surface of the horizontal portion is that of last year's leafy stem. A hemicryptophyte (Fig. 176). Nymphaea {Nuphar) advena.— The cow-lily, or spatterdock, is a hydrophytic cryptophyte with strong horizontal rhizomes, 5 to 10 centimeters thick, and found mostly 8 to 25 centimeters below the soil surface. Where this plant mixes with Scirpus and Typha, it may be classed as a true helophyte, but above it has been classed as a true hydrophyte, the distinction between hydrophytes and helophytes not being defined always sharply. From the rhizome, the surface of which has a check- ered appearance owing to the prominence of the old leaf scars, arise the floating {variegata), or emersed {advena), erect leaves, which are round to ovate, or almost oblong, with an open sinus and subtriangu- lar lobes. The bases of these leaves overlap where they join the rhi- zome. A large number of strong roots grow down from the under side of this rhizome. Figure 175 Hudsonia ericoides. 232 VEGETATION OF THE NEW JERSEY PINE-BARRENS Castalia (Nymphaea) odoraia. — The rhizome of this hydrophyte and cryptophyte is horizontal and much smaller than that of the preceding plant, but found at a similar depth, where a few stout persistent branches may be found. They are 2.2 to 3.2 centimeters thick and 30 centimeters to I meter long, pale in color, covered with a dense, short, black pubes- cence. The apex is rounded, clothed with stipules and fine hairs. The roots, according to Conard, remain uninjured after the evaporation of water from ponds in the fall. In this species 3 to 5 or 6 roots occupy a triangular area on the cushion behind each petiole; the apex of the triangle is farthest from the petiole and is occupied by the first formed root of medium size. Next to it is a second and much larger root, and ^Tt Figure 176 Hypericum densiflorum. Figure 177 Tephrosia virginiana. the base of the triangle has a few smaller ones. A root-cap is present on the tip of every root and large intercellular air-passages in the cortex of the roots. Tephrosia {Cracca) virginiana. — The goat's-rue is a perennial herb with a rootstock that may branch several times. These branches are as thick as a lead-pencil and sink downward obliquely to a depth of 10 centimeters before uniting. The stubs of old aerial roots indicate that some of the rhizome branches have produced at least 5 to 6 annual shoots which are erect and 3 to 6 decimeters tall. The lateral roots, produced 10 centimeters below the surface, are yellowish-white, cord-like, and at least 25 centimeters long. A hemicryptophyte (Fig. 177). VEGETATION OF THE NEW JERSEY PINE-BARRENS 233 3«>' Baptisia tincioria. — The wild-indigo has a strong primary root which descends vertically into the soil and reaches at the soil surface a thick- ness of 2.5 centimeters. In a measured specimen the first lateral root was found 14 centimeters below the soil surface, and it reached a length of I meter, growing downward obliquely into the soil. The shorter lateral roots were found in general at twice that depth, namely, 28 centi- meters. From the crown of this root arise the aerial shoots, usually only one be- ing produced annually, but a number of stubs, at least two, sometimes 4, indicate where former shoots have arisen. A hemicryptophyte (Fig. 178). Lupinus perennis. — The blue lupine is a peren- nial plant with palmately compound leaves of 7-1 1 leaflets. Its stems arise from an under- ground rhizome that may be buried 10 centi- meters under the soil surface. A hemicrypto- phyte (Fig. 179). Galadia regidaris. — The milk-pea is a prostrate perennial with short. Figure 178 Baptisia tinctoria. Figure 179 Lupinus perennis. four- to eight-flowered racemes, from one deeply descending tap- root. Hence its root is of the multicipital kind. A cryptophyte (Fig. 180). Cassia chamaecrista. — This an- nual plant, or therophyte of Raunkiaer's classification, has a slender primary root, about 8 to 10 centimeters long, developed below a subterranean, hypoco- tyledonary portion, 2 centimeters long. The secondary roots, which are produced at intervals along the primary root, bear scattered tubercles. The longest measured lateral root was 22 centimeters long. From six to eight stems may arise Figure i^ Galactia reguiaris. The upright portion of the plant, 234 VEGETATION OF THE NEW JERSEY PINE-BARRENS which is at least 30 centimeters tall, is divided into a number of branches which show at first a plagiotropic tendency, later assuming an upright or orthotropic growth. Hence the branching sys- tem is candelabra-like (Fig. 181). Ilex glabra. — The ink- berry is a shrub 6 to 9 decimeters high, arising from a horizontal root- stock 2 centimeters thick and at least 6 to 7 deci- meters long. Filiform secondary roots arise rather thick A nanophan- FlGURE 181 Cassia chamaecrista from this rootstock. erophyte. Euphorbia ipecacuanhae. — This spurge has a long, vertically directed primary root, which may be over 80 centimeters in length. From the crown of this root, which may be i to 1.5 centimeters thick, arise numerous stems which grow in a diffusely spreading manner. The lateral roots are given off some distance below the surface. One was found at 20 centimeters below and another at 40 centimeters below. A chamaephyte, or a hem- icryptophyte, depend- ing upon whether the origin of the branches is above or below the surface. The radiating branches, which are pros- trate on the ground, bear leaves varying from ob- ovate, or oblong, to narrowly linear, and from a light, glaucous-green to a deep reddish-brown color (Fig. 182). Asclepias amplexicaulis {= A. obtusifolia).— This perennial herb has an upright stem from 3 to 8 decimeters tall, arising from a perpendicular rhizome at least 20 to 30 centimeters in length. Along the rhizome at intervals are borne Figure 183 Asclepias amplexicaulis. Figure 182 Euphorbia ipecacuanhae. VEGETATION OF THE NEW JERSEY PINE-BARRENS 235 opposite lateral buds separated by distinct internodal regions. From the crown of this vertical root- stock may arise two leafy shoots, and a stub is seen frequently Figure 184 Rhus copallina. depth of 7 decimeters to i meter, to grow horizontally at a depth of 6 to 7 centimeters below the surface of the soil. A nanophanerophyte (Fig. 184). Clethra alnifoUa (Sweet Pepper- bush). — This is a shrub which varies greatly in size and grows under a great variety of soil con- ditions. In the pine woods several stems arise from a thick horizontal rhizome, which is provided spar- ingly with secondary roots. A nano- to a meso-phanerophvte (Fig. 185). ^77^ Figure 185 Clethra alnifolia. where the stem of last year was formed. No lateral roots were observed in this plant. A cryp- tophyte (Fig. 183). Rhus copallina. — ^The dwarf sumac is rarely over two meters tall. It arises from a primary root which probably reaches a total The secondary lateral roots begin Figure 186 Chimaphila maculata. 236 VEGETATION OF THE NEW JERSEY PINE-BARRENS Chimaphila macidata (Spotted Wintergreen). — The spotted, leathery leaves of this plant arise from an upright, low stem developed with a rhizome about the thickness of a lead-pencil, buried 5 to 10 centimeters below the surface. A hemicryptophyte (Fig. 186). Dendriiim {Leiophylliim) buxifolium. — The sand-myrtle grows in a tufted manner, many upright, leafy stems arising from a thick, obliquely growing rhizome. A nanophanerophyte (Fig. 187). Figure 187 Dendrium buxifolium. Gaultheria procnmhens. — The wintergreen is a chamaephyte, or hemi- cryptophyte, with evergreen leaves that arise on short, upright branches that are also flower-bearing. These leafy shoots, 5 to 1 5 centimeters tall, are distributed at intervals along the slender, horizontal, creep- ing stems found on, or as much as 5 centimeters below, the surface. It is a dark-brown color. The filiform roots are not over 8 centi- meters long and are found on the under surface of the rootstock. VEGETATION OF THE NEW JERSEY PINE-BARRENS 237 Figure 188 Gaultheria procumbens. A specimen of wintergreen rhizome was measured 48 centimeters long, and another shorter one was 36 centimeters (Fig. 188). Andromeda {N eopieris) mariana. — -The stagger- bush is a nanophanerophyte, 5 to 10 decimeters tall, with oblong to oval leaves and fascicles of nodding flowers in racemes on leafless shoots in the early spring. The upright stems arise from a horizontal rhizome 1.5 centimeters thick, at least 60 to 65 centimeters long. It is found 7 to 8 centimeters below the surface of the soil. The filiform lateral roots are branched and 8 centi- meters long (Figs. 189 and 190). Kalmia angustifolia (Sheep Laurel). — This low evergreen shrub is extremely common in the pine forests. Its leafy stems arise from a much- branched rhizome provided with strong secon- dary roots. A nanophanerophyte (Fig. 191). Kalmia latifoUa. — The laurel is a mesophanero- phyte. Its irregularly shaped, knobby rootstock is chocolate-brown in color and 5 centimeters thick. It grows downward obliquely and joins the rootstock branches of equal size. The lateral roots are much branched, 7 to 10 centi- meters long, and produced most numerously at the crown of the plant (Fig. 192). Epigaea repens. — The trailing-arbutus is a chamaephyte. Its stems, which trail over the ground, arise from a rhizome which merges gradually with the roots, one of which is a tap- root. This is an evergreen plant which it is difficult to grow out of a peaty soil, where its roots, covered with an ectotrophic mycorhiza, find the best conditions Figure 189 Andromeda mariana in flower. Figure 190 Andromeda mariana in fruit. 238 VEGETATION OF THE NEW JERSEY PINE-BARRENS for their development. Coville* has succeeded in growing it from seed (Fig- 193)- Gaylussacia resinosa. — The black huckleberry is a low, much-branched shrub, 0.3 to I decimeter high. It arises from a woody branched rhi- zome with short secondary roots. A nanophanerophyte (Fig. 194). Figure 191 Kalmia angustifolia. Vaccinium pennsylvanicum. — The low, sweet blueberry is character- ized easily by its low habit, bright, warty green twigs, its narrow, lan- ceolate leaves, and its blue-black berries. Its tufted stems arise from a woody rhizome that grows downward obliquely in the humous layer of * Coville, F. V.: Experiments in Blueberry Culture. Bull. 193, Bureau of Plant In- dustry. VEGETATION OF THE NEW JERSEY PINE-BARRENS 239 the soil and produces several strong horizontal secondary roots. A nanophanerophyte (Fig. 195). Vaccinhim vacillans. — This abundant blueberry has a long, creeping rhizome 8 millimeters thick, which is produced 2.5 to 4 centimeters below the soil surface, and grows at least 90 centimeters long. The chocolate-brown lateral roots are tufted and 6 to 8 centimeters long. The upright, aerial stems that arise from this rhizome are 3 to 9 decimeters tall. A nano- phanerophyte (Fig. 196). Chamaedaphne (Cassandra) calyculata. — The leather-leaf is a low, erect shrub with coriaceous leaves that remain for more than a year on the branches and turn reddish-brown in autumn. Dur- ing the early part of the growing season the shoots of the year de- velop large leaves, and toward the end of the growing period small leaves are formed in whose axils are flower-buds that remain closed over winter. Where the tops of the plants project beyond the cover of snow, they may be winter-killed seriously, so that there may be dead leaves and buds at the ex- tremities of the shoots.* Ardostaphylos uva-ursi. — The bearberry is a prostrate, creeping shrub with its branches out- spread on the surface of the ground, but in the course of time the older portions are buried by leaf mold. The last of the Figure 192 Kalmia latifolia. Figure 193 Epigaea repens. year's shoots curve upward slightly, but afterward become more hori- zontal (plagiotropic). The plant grows centrifugally, and its younger * Warming, Eug.: The Structure and Biology of Arctic Flowering Plants, 1908: 32; Gates, Frank C: The Relation of Snow Cover to Winter Killing in Chamaedaphne calyculata. Torreya, 12: 257-262, November, 1912. 240 VEGETATION OF THE NEW JERSEY PINE-BARRENS branches have relatively few adventitious roots, but these secondary roots become more abundant upon the older parts of the branches, especially if covered with soil. The leaves remain green upon the stems from two to three years. The terminal buds on the branches are small in July and they and the lateral buds are protected by the basal part of the petioles of the foliage leaves. The buds bear scale-leaves, greenish or reddish in color, which merge by easy gradations into foliage leaves. In spring the terminal buds are first to develop and the others follow in descending order. The lowermost buds on the year's shoots may remain dormant for several years and may then develop. After the Figure 194 Gaylussacia resinosa. Figure 195 Vaccinium pennsylvanicum. terminal flower-cluster has blossomed, a strong lateral shoot is given off below it, often forming a sympodium with the parent shoot. A chamaephyte (Fig. 197). Pyxidanthera harhulata. — The flowering-moss, or pyxie, is usually a prostrate, or creeping plant, and it shows sun and shade forms. Three different kinds of shoots giving rise to three distinct growth forms may be distinguished in this chamaephyte. First, we have a creeping stem, or rhizome, produced in part under the surface of the soil and in part above the surface. The older parts may be covered with humus to a depth of 1.5 centimeters, and are provided richly with short, brown, filiform roots, VEGETATION OF THE NEW JERSEY PINE-BARRENS 241 while the newer portion is running along on the surface of the soil and is without the filiform rootlets. This form of shoot produces an extended mat- or cushion-plant. Some of the prostrate shoots are at least 20 centimeters long, bearing numerous upright leafy branches which are from 1.5 to 2 centimeters tall and beset with small, linear leaves. Figure 196 Vaccinium vacillans. Figure 197 Arctostaphylos uva-ursi. The second kind of shoot is the periwig type. The shoots are short, and are branched in an upward direction from a common base. They may be covered by humus to a depth of 3 to 4 centimeters, and such buried shoots are covered with brown rootlets. Such a method of branching gives rise to rounded, circumscribed cushions which are usu- ally found in gravel soils or in the bright sunlight. The third kind of shoot occurs on plants that are found in places where there is a consider- able depth of unconsolidated leaf mold, where the shoots, to reach the surface, must fmd their way through the interstices of such superficial deposits. The plant produces in such situations the wander- ing shoots, which bear tufts of loosely aggregated green branches when- ever they reach the surface. The filiform rootlets are found on the older parts of such wandering shoots (Figs. 61, 198, 279, F and G, and design on the front cover of book). 16 =7 "^•^ -s- Tr-~*.A Figure 198 Pyxidanthera barbulata. 242 VEGETATION OF THE NEW JERSEY PINE-BARRENS Figure 199 Melampyrum lineare. Melampynim lineare { = M . americanum) . — The cow-wheat is an erect, branching annual, or therophyte, growing from a primary root which descends obUquely into the soil to a depth of 5 to 6 centimeters, and pro- duces a number of lateral roots which vary in length from 3 to 5 centimeters. It is a common plant in the pine-barrens of New Jersey, growing about 30 centimeters tall, and with opposite, linear, lanceolate leaves (Fig. 199). Chrysopsis mariana. — The golden-aster is a per- ennial herb with silky stems which arise from a weak horizontal rhizome. It grows about i to 3 deci- meters tall. A hemicrypto- phyte (Fig. 202). Hieracium venosum. — The rattlesnake-weed is a perennial herb with an al- most leafless scape, bearing a loose corymb of capitula. The rosette of lanceolate- spatulate, radical leaves arises from the top of a vertical rhizome that is scarcely 2 centimeters long and bears numerous secon- dary roots all over its sur- face. These roots spread in all directions and are at least 10 to 12 centimeters long. The crown of leaves almost meets the upper roots, so that not over 5 millimeters of actual rhizome surface is bare either of roots or attached leaf bases. A hemicryptophyte (Fig. 200). Sericocarpus asteroides ( = 5. conyioides). — The white-topped aster is a perennial herb, 35 centi- meters tall, and leafy to the top. The under- ground parts of this plant are not over 2 centi- meters below the surface of the soil. There is a short horizontal rhizome that is branched usually, from which arises an intricate mass of strong secondary roots, some of Figure 200 Hieracium venosum. Figure 201 Eupatorium verbenas- folium. VEGETATION OF THE NEW JERSEY PINE-BARRENS Figure 202 Chrysopsis mariana. Figure 203 Sericocarpus asteroides. 244 VEGETATION OF THE NEW JERSEY PINE-BARRENS them 1 5 to 20 centimeters long, growing 2 centimeters below the surface. Usually only one upright stem is developed from a single branch of the rhizome. I n addition the plant develops a series of suckers. One plant in the field had six of these suckers, bearing pinkish colored scale-leaves with adventitious sec- ondary roots along their course of 8 centimeters, and a tuft of petiolate, spatulate leaves at their extremity. Such a plant may be described as surculose, and according to Raun- kiaer's classification, it is a hemi- cryptophyte (Fig. 203). Eupatorium verbenaefolium. — This plant is an erect cryptophyte with aerial stems 50 centimeters tall, bearing opposite, ovate-lance- olate, coarsely toothed, sessile leaves. This upright stem arises from a corm-like base, buried 5 centimeters below the soil surface. Numerous roots arise from this rounded base, and the longest one measured was 30 centimeters long, and it grew down obliquely into the soil. Themajority of the upper roots grew almost horizontally (Fig. 201). Liatris graminifolia. — The but- ton-snakeroot is a hemicryptophyte with an upright stem 3 to 9 deci- meters tall, arising from a corm-like base from i to 2 centimeters in di- ameter. Numerous white, filiform roots 3 to 4 centimeters long arise from its under surface, and two to several upright leaf- and flower- bearing stems from its upper side about 4 centimeters below the soil Figure 204 surface. Several buds arise at the Liatris graminifolia. top of the COrm (Fig. 204). CHAPTER XVII GENERAL REMARKS ON ROOT DISTRIBUTION The distribution of the roots and underground parts of pine-barren species is determined by several important conditions which must be considered in any detailed account of the ecology of pine-barren vegeta- tion. The first important factor is the living plant considered specific- ally, as to the structural characteristics of its underground system. Each plant has specific root and underground stem characters, which are ancestral, and which are subject only to differences of length, thickness, color, and other minor modifications due to the environment (Fig. 205). The second important factor in considering the subject of the distribu- tion of the underground parts of pine-barren plants is the actual associa- tion of the species together where the growth of one kind of root system excludes the growth of another kind. Thirdly, the soil structure, or texture, has great influence in determining the distribution of the under- ground systems of these plants, as also the ground water, the tempera- ture of the soil and its aeration. The bacterial and fungous floras of such soils also have an important bearing upon the distribution through the soil of pine-barren roots and root-stocks. In the preceding pages we have described the subterranean organs of different species of pine-barren plants, and in this study we have empha- sized the specific differences of structure and growth in the soil. The soil structure, or texture, its permeability or impermeability, its color and other peculiarities, have been discussed in detail, but the following notes, taken in the field, may be inserted at this point as directly appli- cable to the discussion. The following is a series of measurements of the strata of pine-barren soils taken at a number of widely separated locali- ties in the region: Pine Forest, Como Lake: Leaf Mould Layer 6 centimeters White Sand Layer 27 Red Gravel Layer 33 Pine Forest, Avon: Upper Humous Layer 16 White Sand Layer 30 Red Gravel Layer 29 245 246 VEGETATION OF THE NEW JERSEY PINE-BARRENS Pine Forest, Shamong: Upper Humous Layer 8 centimeters White Sand | Root-penetrated 16 cm. | I Rootless 16 cm. J Red Sand Layer Unknown depth* Pine Forest, Sumner: Upper Humous Layer 20 centimeters White Sand Layer 40 Reddish Sand Layer Unknown depth Upper Plain: Upper Humous Layer 15 centimeters White Sand Layer 30 Yellow Sand with Limonite 1.2 meters Light Yellow Sand with Ground-water. Deciduous Forest, Clementon: Upper Humous Layer 30 centimeters Middle Soil Layer 25 Water Table We have dfecussed the successive layers found in a pine-barren forest, beginning with the tall pine trees, then the undergrowth of oaks, then the shrubs, then the undershrubs, the herbaceous plants, and those prostrate on the surface of the soil. Similarly, we find strata of the under- ground parts of the plants, which ramify through the layers of the soil above described. We wish to ascertain now the layer of the soil where most of the roots and rhizomes occur and after we have definitely located them to draw conclusions therefrom. Of these 55 species, 4 are aquatic, 5 are bog plants, leaving 46 as true pine forest species. Of this 46, we find that 5, or one-ninth, are surface growers, or chamaephytes, such as Gaultheria procumbens, Epigaea repens, Arctostaphylos uva-ursi, and the flowering-moss, Pyxidanthera barbulata (Fig. 198). The demands then that these plants make on the soil is only on the most superficial part of the upper humous layer. Twenty-six species occupy the hemi- cryptophyte layer and their roots and root-stocks live in part in the upper humous layer and in the topmost part of the white sandy layer. Such are arranged systematically as follows: Andropogon scoparius, Xerophyllum asphodeloides, Smilax glauca, Cypripedium acaule, Comp- tonia asplenifolia, Helianthemum canadense, Hudsonia ericoides, Are- naria caroliniana, Hypericum densiflorum, Comandra umbellata, Teph- rosia (Cracca) virginiana. Cassia chamaecrista, Andromeda mariana, Vaccinium vacillans, V. pennsylvanicum, Gaylussacia resinosa, Kalmia * At Lakewood the red-gravel layer of virgin soil in a gravel pit was found to be 9.1 meters deep, and in a railroad cut, 4.5 to 6 meters thick. a, Panicum dcpauperatum. aa, Andromeda (Pieris) mariana. b, Baptisia linctoria. bb, Vaccinium pi-nnsylvanicum. c, Cypripediiini acaiilo. cc, Pyxidanthera barbulata. Figure 205 Diagrammatic reprosontalion of the atrial and root systems of the plants of the pine-barren formation. (The horizontal lines below the soil surface represent successive soil strata. See pages 245 and 246.) d, Gaylussacia resinosa. ff, Cladonia rangiferina. ii, Quercus stellata. II, Andropogon scoparius. r, Melampyrum americanum. dd, Quercus marylandica. g, Sericocarpus asteroides. j, Chimaphila maculata. m, Quercus prinoides. s, ddd, Hypericum densillorum. gg, Smilax glauca. jj, Vaccinium vacillans. n, Lupinus perennis. t, c, Andropogon scoparius. h, Kalmia angustifolia. k, Asclepias obtusifolia. o, Dendrium buxifolium. u, ec, l-'pigaea repens. hh, Xerophyllum asphodeloides. kk, Liatris graminifolia pilosa. p, Tephrosia (Cracca) virginiana. Kalmia latifolia. Arctostaphylos uva-ursi. Gaultheria procumbens. Sassafras variifolium. X, Fvuphorbia ipccacuanhae. y, (;lelhra ainifolia. z, Mieracium venosum. P, Pinus rigida. f, (Juercus ilicifolia. Comptonia asplenifolia. I, Hudsonia ericoides. q, Eupatorium verbenaefolium. w. Cassia chamaecrista. VEGETATION OF THE NEW JERSEY PINE-BARRENS 247 angustifolia, Dendrium buxifolium, Melampyrum lineare, Eupatorium verbenaefolium, Liatris graminifolia, Chrysopsis falcata, Solidago odora, Aster spectabilis and Hieracium venosum. The next deeper layer of soil, which comprises practically the white sandy layer and the topmost part of the reddish sand, or gravel beneath, is occupied by the roots and rhizomes of Pteris aquilina, horizontal roots of Pinus rigida, Myrica carolinensis, Quercus prinoides, O. marylandica, Q. ilicifolia, Q. stellata, Sassafras variifolium, Baptisia tinctoria, Ilex glabra. Euphorbia ipecacuanhae, Asclepias amplexicaulis, Rhus copal- lina and Kalmia latifolia. In some cases the upper lateral roots of these 14 pine-barren species are found in the upper humous layer, entering into competition with the complex of roots of the plants listed above. The reddish sand and gravel layer beneath the sandy layer is penetrated by few roots, and the roots that penetrate are deep-growing, primary roots of the principal trees, such as: Pinus rigida, Quercus marylandica, Q. stellata, and such shrubs as Q. ilicifolia, Q. prinoides, Kalmia latifolia, and Rhus copallina. From this enumeration it is clear that the roots of the majority of pine- barren plants are superficial ones found in the first 30 centimeters of the surface soil, which would include the compact humous layer below the loose leaf mould layer, which need not be included, and the white sandy layer, which includes a considerable amount of humus, and therefore ought to be included strictly as part of the humous layer (Fig. 205). The roots and root-stocks of the perennial plants of the pine-barrens make a continuous demand upon the soil for water and food substances. The annual plants, or therophytes, such as Melampyrum lineare. Cassia chamaecrista, and others absorb materials from the soil only during the summer months, when they make a rapid growth. The same classifi- cation of root systems can be made for pine-barren species that Cannon has made for the desert plants of the southwest: (i) Root-systems, which extend horizontally from the main plant and lie for their whole course near the surface of the ground; (2) root systems which are characterized by a strongly developed tap-root going down directly to a depth determined in part by the character of the soil, in part by the penetration of rain-water, and in part by the character of the root itself; (3) and roots that reach not only widely, but penetrate also fairly deeply. The majority of the pine-barren plants rooted in the humus and with horizontal subterranean organs belong to the first category. The pines belong to the second category and the oaks to the third (Fig. 205). With these facts in mind the distribution of the rainfall throughout 248 VEGETATION OF THE NEW JERSEY PINE-BARRENS the year must be considered in relation to the growth and successful competition of the various pine-barren species of plants. It is important to emphasize the statement of Kraus* that, "under natural conditions, one may fmd in the smallest areas an endless diversity of chemically and physically distinct habitats," for there can be no criticism, if one cites the general conditions that prevail in the region. An estimate of the amount of rain and melted snow throughout the year expressed in inchest gives the following for two localities in the pine-barrens. January 3 February 3 March 4 April 2 May 2 June 4 July. August 5 September 4 October 2 November 3 December 3 IN ELAND 4.60 4 06 4 43 3 12 3 76 3 52 4 25 5 09 4 3H 3 33 3 72 4 01 Total 4676 48.27 But that this precipitation is not always uniformly distributed in the months of the year is indicated from the table of extreme monthly precipitation in inches and hundredths for Vineland: Greatest January 6.30 February 6.25 March 6.84 April . May. 8.32 8.45 June 5-59 July 982 August • • • 10.64 September 9-9i October 6.75 November 7-24 December 7-52 Lowest 1.30 1-73 1.22 ••73 0.77 0.60 1.85 1.28 0.69 1.08 1.49 1.88 At Vineland, in 1881, the rainfall for July, August, and September amounted to 6 inches, as compared with the average of 14.2 inches. And there were two periods of twenty-one and twenty-two days re- spectively when no rain fell. As a general thing it may be stated that, with ordinary rains, the rain-water does not penetrate the soil to any * Kraus, G.: Boden und Klima auf kleinstem Raum. Versuch einer exakter Behand- lung des Standorts auf den Wellenkalk, pp. 184, pis. 7, Jena, 191 1; Reviewed in The Plant World, 15: 300, December, 1912. t All the United States Weather Bureau records are in inches. VEGETATION OF THE NEW JERSEY PINE-BARRENS 249 considerable depth, first, because there is not enough of it, and second, the thirsty roots of the first 30 centimeters (1 foot) of sandy humous soil absorb it and very little percolates into the lower soil layers. The water-table lies usually so deep that it is not available as a source of water supply to the superficially rooted plants, for the capillary rise of water is extremely slow.* With heavy rains, or a steady one of several days' duration, the rain-water drains into the subsoil, which is replenished thereby. I have no meteorologic records which give the number of such rains throughout the year, but they are few and far between. The length of time that the water in available amounts remains in the soils is a vari- able one. But from the first table above it is seen that the rainfall is suf- ficient during the year for the superficially rooted annuals and perennials which can utilize all of the water, which falls as rain, whether the showers be heavy or light. But there are critical periods when no rain falls and then the xeroph>tic structures of such plants as Kalmia latifolia, Gaultheria procumbens, Epigaea repens, Xerophyllum asphodeloides, Ilex glabra, Arctostaphylos uva-ursi, tide them over the period of drought. With the deep-rooted trees, it is otherwise, for during the critical period of dry weather and the cold dry period of winter, their deep vertical and superficial horizontal root systems enable them to get a supply of water under all climatic conditions quite sufficient to supply the loss by transpiration. During dry weather the deeper roots are depended upon to secure the water lost in transpiration; during wet weather, or when the surface soil is wet, the superficial roots are active. The amount of transpiration is under control in our northern pinesf in which the surface exposed by the evergreen, acicular leaves, as a whole, renders it necessary for the individual leaves to be xeromorphic and xerophytic. This structure enables these coniferous trees to live in regions where there is a season of physiologic drought. The tracheidal structure of their wood is well suited to their xerophytic evergreen leaves. With the horizontal root system the species of pine growing in the New Jersey pine-barrens can secure additional supplies of water from the surface layers after a light shower. The deciduous trees, as trophophytes, are potential xerophytes, for in winter with the fall of their leaves the rate of transpiration is materially reduced. As the pine-barren soils have a fair capacity for retaining water and occupy an intermediate position with respect to the rate of percolation of water * Ante, under description of soils. t Groom, Percy: Remarks on the Oecology of Coniferae. Annals of Botany, xxiv, 241-269, April, 19 10. 250 VEGETATION OF THE NEW JERSEY PINE-BARRENS through them, there is probably no period of the ordinary year when some water is not available to the native plants of the region.* The pine-barren soils, being open and pervious to water, the sunlight being intense and the sand at times heated highly by the sun's rays, the plants that live under such conditions must have been modified struc- turally by the influence of their environment, so as to become drought resistant. This implies adaptations for enduring drought, adaptations for evading drought and adaptations for escaping drought. In the presence of water-storage tissues, as in succulent roots, stems and leaves and in the ability to become dormant during the critical season of the year pine-barren plants show adaptations for enduring drought. The various structural peculiarities by which transpiration is controlled, or by which the root system has an excessive development are adaptations for evading drought. Finally, when a plant is annual and grows only during the favorable season and then dies to the ground, having liberated its seeds, it escapes the period of cold, or physiologic dryness. In the pine-barrens the first and third methods of resisting drought are rather infrequent, but structural peculiarities of leaf and exceptional root development are not uncommon. Another matter of interest should be mentioned at this juncture. The conductive power of the pine-barren soils is such that they become heated intensely by the sun's rays falling directly upon them. In short, they are warm soils. They often become so hot in the heat of the day as to be unbearable to the touch of the bare foot or hand. The influence of such sandy soil is both heating and drying, and somewhat like that of a desert, making the days hot and the nights, owing to the rapid radia- tion of heat, cool. These factors have a decided influence on the char- acter of the vegetation of the region. Having determined the depth at which the subterranean organs of pine-barren plants are found, we can appreciate why this type of vegetation of superficially rooted perennials has been able to occupy southern New Jersey, as a climax formation, since very early times (see ante). The matted and interlaced condition of the roots and root-stocks of the herbs, shrubs and trees is so intricate that no alien plant has a chance of establishing itself in an area where the original vegetal covering has remained unbroken. There is such an interrelation and reciprocity of underground parts that each is found at the soil level where its functions can be performed best and where there is soil space enough to grow. This adjustment of the root systems of the pine-barren plants which grow in association has taken ages to bring to its present state, and such plants as enter into the different formations * See ante under Soils. VEGETATION OF THE NEW JERSEY PINE-BARRENS 25 I grow together because their root systems are in a complicated reciprocity of growth, so that the soil space is filled after a long period of competition. Such competition by which many species were replaced by others has led to an adjustment in which, instead of competition, the root organs each fill a particular soil space. Through the survival of the fittest the struggle for existence among the different native species has become an almost negligible factor. Such plants naturally exclude other species that have subterranean parts that are unable to be adjusted to the com- plex of the roots of the long-established pine-barren plants. The moisture conditions under which a species grows may be those which prevent the encroachment of competing species. The non-entry of some species is determined probably during the early stages of germi- nation and growth of the seedling. Obviously, says Yapp,* if the con- ditions are such as are very unfavorable to the young growth, there is a poor chance of its surviving to the adult stage. (Compare Fig. 205.) With the few water plants to which we have drawn attention it is otherwise. During the entire year there is water sufficient for their normal growth, and hence, although we have classed them as crypto- phytes, yet it is mainly with reference to the depth below the surface of the muddy bottom of the ponds and the streams where their roots and root-stocks are found. Such plants as Tofieldia racemosa, Pogonia ophioglossoides, Calopogon pulchellus, Habenaria blephariglottis, and the shrub, Chamaedaphne calyculata, have usually sufficient water throughout the year, except in cases of unusual drought, when the sphagnum dries up. * Yapp, R. H. : Sketches of Vegetation at Home and Abroad, iv. Wicken Fen, The New Phytologist, VII : 76. CHAPTER XVI II LEAF EORMS OF PINE-BARREN PLANTS The older text-books of botany, for the sake of training the botanic stu- dent in the identification of species by the use of manuals, or floras, em- phasized the descriptive terms used in describing the shapes, margins, apices, and other parts of leaves. These terms are still in common use, but latterly considerable emphasis has been given to the study of leaf forms from the ecologic and morphogenetic points of view. Goebel was one of the first of modern botanists to break the leashes of the older morphology. In his classic Pflanzenbiologische Schilderungen (1889- 1893) he considers the form of the leaf from its morphogenesis. Experi- mental methods are used, which are nowhere better illustrated than in his discussion of water plants, where the influence of land conditions and water conditions are determined experimentally for a number of water plants. Sir John Lubbock, in his little book, "Flowers, Fruits and Leaves" (1894), approaches the subject from much the same stand- point, and Henslow, in his illuminating book, the "Origin of Plant Struc- tures" (1895), gives in Chapter XII a modern statement of the form and structure of leaves. L. H. Bailey devotes 52 pages of "Lessons with Plants" (1898) to a study of leaves and foliage. Warming (1901), in a brochure entitled "Om Lovbladformer,"* gives an interesting resume of the subject. Sir John Lubbock gives a chapter to leaves in his " Notes on the Life History of British Flowering Plants" (1905), and Goebel, in Part II of his monumental work, the "Organography of Plants," presents in the English translation of his work (1905) a full synopsis of the views of modern morphologists on the origin, shape, and structure of various leaf forms. If we analyze the leaf forms of the 555 native pine- barren species, our analysis shows that the following leaf forms exist in the region. As a number of duplications occur in the following list, and as it was found difficult to classify exactly some of the leaf forms, the total number of leaves listed in the following arrangement does not total the entire number of 555 pine-barren species of plants. This discrep- * Warming, EuG.: Oversigt over det Kgl. Danske. Videnskabernes Selskabs Forhand- linger, 1901, No. i. 252 VEGETATION OF THE NEW JERSEY PINE-BARRENS 253 ancy does not detract, however, from the value of the enumeration of all the different kinds of leaf forms. In the groups the species are arranged systematically. 1. Leaves with Circinate Ptyxis, Some as Sporophylls. — The leaves of this group show a blade that is characterized by apical growth and a manner of unfolding that is called circinate. The group includes the ferns: Osmunda regalis, O. cinnamomea, Schizaea pusilla, Lygodium palmatum, Pteris (Pteridium) aquilinum, Woodwardia virginica, W. areolata, Asplenium platyneuron, A. filix-foemina, Polystichum acros- tichoides, Dryopteris thelypteris, D. simulata, Phegopteris dryopteris. = 13- 2. Lycopodium, or Cedar Form. — This type is exhibited in the club- moss, genus Lycopodium, with an axis beset with numerous small lanceolate or subulate leaves, arranged in two to many ranks. In the pine-barrens this type is represented by four species of Lycopodium, and by the trees Chamaecyparis thyoides, Juniperus virginiana, and the herbs, Arenaria caroliniana, Pyxidanthera barbulata. =8. 3. Needle Leaves in Fascicles. — The acicular leaves of the pines are found in fascicles of two in Pinus echinata and in threes in Pinus rigida. They are evergreen. =2. 4. Algoid Leaves. — These are narrow submerged leaves which re- semble the thallus of certain algae in their long and narrow blades. Such leaves are found in Potamogeton confervoides and Zannichellia palustris. = 2. 5. Linear, or Grass Form. — This group includes 142 species of grasses and sedges. Although the sedges do not always have the gen- eralized type of linear leaf, yet it is unnecessary for our purpose to make a separation of this form into a number of different categories. Here we may include the leaves of Schizaea pusilla (a fern), Sagittaria graminea, 5 species of Xyris, 4 species of Eriocaulon, 12 species of Juncus, Tofieldia racemosa, Abama americana, Xerophyllum asphodeloides, Zygadenus leimanthoides, Gyrotheca tinctoria, Hypoxis hirsuta, Iris prismatica, Sisyrinchium angustifolium, S. atlanticum, Euthamia graminifolia (radical leaves), E. tenuifolia (radical leaves), Sericocarpus linifolius (radical leaves). =177- 6. Heather Form. — The leaves are small and closely crowded on the shoots of plants of this type. Here may be mentioned Corema Conradii, Hudsonia ericoides, H. tomentosa. =3. 7. Minute Subulate Leaves. — These leaves are small, but they are scattered on the stem, otherwise types 6 and 7 might have been com- bined. The following plants have small leaves: Sarothra gentianoides, 254 VEGETATION OF THE NEW JERSEY PINE-BARRENS Lechea Leggettii, Bartonia virginica, B. paniculata, Utricularia cornuta, U. juncea, U. subulata, U. cleistogama. =8. 8. Linear Leaves, not Grass-like. — The linear leaves of the fol- lowing plants are usually cauline and without overlapping bases and sheaths, as in grasses and sedges: Potamogeton epihydrus (submerged leaves), Polygonella articulata, Linum medium, L. floridanum, Polygala viridescens, P. mariana, P. Nuttallii, Crotonopsis linearis, Linaria canadensis, Ludwigia linearis, Kneiffia linearis, K. longipedicellata, Trichostema lineare, Stachys hyssopifolia, Diodia teres, Lobelia Nut- tallii, L. Canbyi, Sclerolepis uniflora, Lacinaria graminifolia, Euthamia graminifolia Nuttallii, E. tenuifolia, Sericocarpus linifolius, lonactis linariifolius, Helianthus angustifolius, Coreopsis rosea, Chrysopsis fal- cata, Anaphalis margaritacea, Gnaphalium obtusifolius. The specific names of a considerable number of these plants refer to the linear leaf form. =28. 9. Iris, or Typha Leaf Form. — The erect, sword-shaped blade of leaves of this type is familiar in the Iris, or blue-flag. Relatively few pine-barren plants show this type and all that have this form of leaf are aquatic: Acorus calamus, Typha latifolia, Sparganium americanum, S. americanum androcladum, and Iris versicolor. 10. Simple Lanceolate Leaves (Willow and Knot-weed Type). — This group includes a diverse number of plants of no close systematic affinity, which display a lance-shaped stem leaf. For convenience we will divide them into trees, shrubs, and herbaceous plants. (a) Trees and Shrubs. — Salix nigra, S. humilis, S. tristis, Myrica caro- linensis, Comptonia asplenifolia, Quercus phellos, Itea virginica, Spiraea latifolia, S. tomentosa, Malus angustifolia, Aronia arbutifolia, Amelan- chier intermedia, Prunus serotina. Ilex verticillata, 1. laevigata, llicioides mucronata, Nyssa sylvatica. Viburnum nudum. =18. (b) Herbs. — Polygonum emersum, P. Careyi, Crotalaria sagittalis, Acalypha gracilens, Ludvigia alternifolia, Chamaenerion angustifolium, Epilobium coloratum, Asclepias tuberosa, Myosotis laxa, Onosmodium virginicum, Verbena urticifolia, V. angustifolia, Lycopus virginica, L. sessilifolius. Lobelia inflata, Solidago erecta, S. puberula, S. semper- virens, S. odora, S. rugosa, S. fistulosa, S. neglecta, S. uniligulata, S. nemoralis, Aster novi-Belgii, A. nemoralis, A. dumosa, Helianthus divaricatus and Bidens connata. =29. 1 1 . Yucca Form. — This type is a sword-like blade, which is somewhat leathery in texture. It is represented by one pine-barren plant, viz., Eryngium yuccifolium. 12. Radical Lanceolate or Spatulate Leaves. — There are quite VEGETATION OF THE NEW JERSEY PINE-BARRENS 255 a few plants with clustered leaves arising as a rosette from the crown of the root or root-stock. Besides the radical leaves, there are cauline ones. Here are included: Adopogon virginicum, Hieracium venosum, H. Gronovii, Solidago erecta, S. puberula, S. stricta, S. sempervirens, S. rugosa, S. neglecta, S. uniligulata, Sericocarpus asteroides, Aster spectabilis, A. gracilis, Antennaria neglecta, Gnaphalium purpureum, Senecio tomentosus. = i6. 13. Simple Broad Deciduous Leaves. — (a) Unlohed. — Populus grandidentata, Betula populifolia, Alnus rugosa. {h) Lohed. — Quercus coccinea, Q. velutina, Q. digitata, Q. ilicifolia, Q. marylandica, Q. alba, Q. stellata, Q. prinus, Sassafras variifolium, Acer rubrum carolinianum, Vitis aestivalis. = 14. 14. Bay-like Leaves. — These leaves are broad with almost entire margins, evergreen and leathery, resembling those of the true laurel, Laurus nobilis. The group includes: Smilax laurifolia. Magnolia glauca, Kalmia latifolia, Rhododendron maximum, Ilex opaca, I. glabra. =6. 15. Smilax-Ipomoea Leaf Form. — The leaves of these two genera of climbing plants are broad approaching heart-shaped. The group in- cludes Smilax tamnifolia, S. rotundifolia, S. glauca, S. Walteri, Dioscorea villosa, Ipomoea pandurata, Convolvulus sepium. =7. 16. Violet Leaf Form. — This group was made to include the pine- barren species of the genus Viola, such as: Viola pedata, V. sagittata, V. fimbriatula, V. emarginata, V. primulifolia, V. lanceolata. =6. 17. Broad Sagittate Leaves. — Only two plants have typic arrow- shaped leaves, viz., Sagittaria longirostra and Nymphaea variegata. 18. Dandelion Form. — The leaves of the dandelion, Taraxacum officinale, are incised-pinnatifid. A few pine-barren plants have leaves of this type: Oenothera laciniata, Dasystoma pedicularia, D. flava, D. vir- ginica, Adopogon carolinianum, Lactuca canadensis, L. hirsuta. =7. 19. Nabalus Leaf Form. — The genus Nabalus, or Prenanthes, in- cludes herbs with alternate, mostly petioled, dentate, lobed, or pinnatifid leaves. The upper are sometimes auriculate and clasping. In the pine- barrens of New Jersey there are two species, Nabalus serpentarius and N. virgatus. =2. 20. Leathery Leaf. — This type of leaf may be described as coria- ceous or ericaceous, because common in that family. {a) Deciduous. — Clethra alnifolia, Azalea nudiflora, A. viscosa, Leu- cothoe racemosa, Andromeda (Neopieris) mariana, Xolisma ligustrina, Gaylussacia frondosa, G. dumosa, G. baccata, Vaccinium corymbosum, V. virgatum, V. caesariense, V. atrococcum, V. pennsylvanicum, V. vacillans. =15. 256 VEGETATION OF THE NEW JERSEY PINE-BARRENS (b) Evergreen. — Pyrola americana, P. elliptica, P. chlorantha, Chima- phila maculata, C. umbellata, Dendrium buxifolium, Kalmia angusti- folia, K. latifolia, Epigaea repens, Chamaedaphne calyculata, Gaul- theria procumbens, Arctostaphylos uva-ursi, Vaccinium macrocarpon. = •3- 21 . Compound Pinnate Leaves. — This heading is a term of the older morphology. Warming has emphasized the importance of enumerating the plants that have simple and compound leaves in any particular geographic region, as that of the pine-barrens of New Jersey. The following plants have compound pinnate leaves: Hicoria alba, H. glabra, Drymocallis arguta, Rosa humilis, Cassia nictitans, Tephrosia (Cracca) virginiana, Apios tuberosa, Rhus copallina, R. vernix, Aralia nudicaulis, A. spinosa, Oxypolis rigidior. =12. 22. Compound Trifoliate Leaves. — The compound leaves with three leaflets are more common in the pine-barrens than the preceding type. The plants with trifoliate leaves are: Rubus cuneifolius, R. argutus, R. villosus, R. hispidus, Baptisia tinctoria, Stylosanthes biflora, Meibomia Michauxii, M. sessilifolia, M. stricta, M. viridiflora, M. Dil- lenii, M. rigida, M. marylandica, M. obtusa, Lespedeza repens, L. frutescens, L. virginica, L. hirta, L. oblongifolia, L. capitata, L. angusti- folia, Clitoria mariana, Galactia regularis and Rhus radicans. =24. 23. Compound Palmate Leaves. — A single plant has leaves of this type, viz., Lupinus perennis. =1. 24. Variable Leaf Forms. — Euphorbia ipecacuanhae belongs to this group. The wild-ipecac or ipecac spurge, has leaves that vary from green to red and in outline from linear to orbicular. It is a remarkable plant, because these leaf forms are found on plants that grow in close proximity to each other. = i . 25. Floating Leaf Forms. — The water plants with floating leaves are Brasenia purpurea, Nymphaea variegata, Castalia odorata, Lim- nanthemum lacunosum, Utricularia inflata, Potamogeton Oakesianus, P. epihydrus, Orontium aquaticum. =8. 26. Dissected Submerged Leaves. — In aquatic plants the sub- merged leaves have frequently a different conformation from those which stand above the surface of the water, or which float upon it. The influence of water, illumination, and temperature, is to produce a di- vided or dissected leaf and in some types a ribbon-shaped leaf is found. The pine-barren plants with dissected leaves are Proserpinaca pectinata, Myriophyllum humile, M. heterophyllum, Utricularia intermedia, U. inflata, U. clandestina, U. purpurea, U. gibba, U. fibrosa, Potamogeton Oakesianus. =10. VEGETATION OF THE NEW JERSEY PINE-BARRENS 257 27. MoNOTROPA Leaf Form. — As the Indian-pipe, Monotropa uni- flora, is a saprophyte, the reduced leaves of this and other plants are related to a saprophytic or a parasitic life. The leaves of the following plants are colorless, scale-like and closely appressed: Monotropa uni- flora, Monotropa hypopitys. =2. 28. Dodder Form. — The dodders (Cuscuta) are parasitic plants attached by haustorial roots to various host plants. The leaves, as a consequence of this parasitic life, are reduced to a few scale leaves around the clusters of flowers. Here belong Cuscuta arvensis, C. com- pacta. 29. Fly-trap Leaf Form. — ^The leaves of the Droseras vary from linear to orbicular. The surface of the blade is covered with gland-bear- ing filaments or tentacles. Each gland is surrounded by a drop of viscid secretion, which glistens in the sunlight, hence the name, sundew. If a small insect alights on the leaf, it is caught by the viscid secretion. The tentacles bend over the insect until it is firmly held and digestion proceeds until the soft parts of the insect are dissolved and absorbed. In the pine-barrens there are three species: Drosera rotundifolia, D. longifolia, D. filiformis. =3. 30. Pitchered Leaf Form. — The ascidial leaves of the pitcher-plant, Sarracenia purpurea, are well known to catch insects, but it is doubtful whether any digestion takes place. It is more probable that the pitcher filled with insects decays and the putrid organic material, in part the dead bodies of insects, in part plant tissue, fertilizes the soil in which the plant grows. = i. 31. Bladder Leaf Form. — The insect-trapping bladders of Utricu- laria are modified leaves. Not all of the pine-barren species have these bladders. The following five species are provided with them: Utricu- laria fibrosa, U. clandestina, U. gibba, U. inflata, U. purpurea. =5. 32. Clasping Leaves. — Two plants have leaves attached to the stem in this manner: Specularia perfoliata and Aster patens. =2. 33. Leaves Opposite (Herbaceous). — The following plants have opposite leaves: Silene antirrhina, Sagina decumbens, Arenaria caro- liniana, Acer rubrum caroliniana, Ascyrum stans, A. hypericoides, Hypericum densiflorum, H. maculatum, H. boreale, Triadenum virgini- anum, Elatine americana, Rhexia virginica, R. aristosa, R. mariana, Isnardia palustris, Lysimachia terrestris, Sabatia lanceolata, Gentiana porphyrio, Apocynum androsaemifolium, A. cannabinum pubescens, Asclepias rubra, A. variegata, Verbena urticifolia, V. angustifolia, 1 1 species of the family Labiatae, Veronica peregrina, Dasystoma pedicu- laria, D. flava, D. virginica, Oldenlandia uniflora, Cephalanthus occi- 17 258 VEGETATION OF THE NEW JERSEY PINE-BARRENS dentalis, Mitchella repens, Diodia teres, Viburnum cassinoides, V. nu- dum, 7 species of Eupatorium, Coreopsis rosea and Bidens connata. = 54. 34. Verticillate Leaves. — The following are some of the pine-bar- ren plants with verticillate leaves: Lilium superbum, Medeola virginica, Polygala cruciata, P. brevifolia, P. verticillata, P. ambigua, Decodon verticillatus, Lysimachia quadrifolia, Trientalis borealis, Asclepias ver- ticillata, Monarda punctata (bracts included), Galium pilosum, Eupa- torium maculatum, E. hyssopifolium. =14. General Remarks on Leaf Forms We have enumerated in the preceding 34 groups 518 species of plants, some of which have been included twice and occasionally 3 times. However, this can be said, that the whole pine-barren flora of 555 native species has been pretty well covered in the above classification. Only those forms of leaves have been omitted which have been considered relatively unimportant or which it has been impossible to classify satis- factorily. In order to compare the leaf forms of the pine-barren region with two other phytogeographic regions, we will divide all pine-barren plants into two groups: those with simple leaves and those with com- pound leaves. By making this distinction, we can correlate the figures thus obtained with those given by Warming for Denmark and Lagoa Santa in South America. Of the 555 native pine-barren species, 50 species have compound leaves and 505 have simple leaves. The list of 50 species with compound leaves includes 1 3 ferns with pinnately com- pound fronds. Excluding the ferns, there are 37 species with compound leaves and 542 species, less 37 species, or 505 species with simple leaves, or as I : 13.6. Warming estimated that there were 410 species of Danish vascular plants comprising the forest flora of the country and 1000 species in the open country. In the woods of Denmark there exist approximately 180 species with leaves divided or compound, and 230 species with simple leaves. The ratio is then as 3 : 4. In the open places, he estimated that there were 750 species with simple leaves and 250 species with divided or compound leaves. The proportion is 3 : i . Warming, in his work, " Lagoa Santa," which deals with a portion of the Brazilian vegetation, enumerates 300 trees and herbaceous or woody plants, and the larger majority of these plants have simple leaves (lance- olate, elliptic, etc.). CHAPTER XIX MICROSCOPIC LEAF STRUCTURE In two previous papers* I have described the leaf structure of the sand- dune plants of Bermuda and of the strand plants of New Jersey, namely, the dune plants and the salt-marsh plants. Bibliographic notes are given in each of these papers, so it would be superfluous to repeat what 1 have emphasized. Since these papers were written a number of recent accounts have been published and these bear directly upon the micro- scopic leaf structure of plants. Woodhead,! in his " Ecology of Wood- land Plants in the Neighborhood of Huddersfield," gives an excellent account of the effect of environment on structure, using 8 species as examples. The structure of each species is illustrated by figures of the microscopic structure of stems and leaves. Later, in 1910, appeared the magnificent monograph of Jean Massart entitled, " Esquisse de la Geo- graphic Botanique de la Belgique." Considerable attention is given in this book to the leaf structure of a selected number of Belgian plants of ecologic interest. The October (1912) number of the Botanical Gazettel contains an illustrated study of the "Comparative Anatomy of Dune Plants," by Anna M. Starr. Only two of the plants studied by her are found in the pine-barren region of New Jersey. The response of leaf structure to light and transpiration is found in the palisade layers, their presence on the upper and under sides of the leaves and their arrangement, so that the central part of the leaf be- comes palisade throughout, if the light is intense and the transpiration excessive. When both leaf surfaces are illuminated equally, the leaf may be termed isophotic; when illuminated unequally, diphotic. Di- photic leaves which show a division into palisade and spongy parenchyma have been called by Clements diphotophylls. Isophotic leaves are of three types, viz., the staurophyll, or palisade leaf; the diplophyll, or double leaf; the spongophyll, where the rounded parenchyma cells * Harshberger, J. W.: The Comparative Structure of the Sand-Dune Plants of Ber- muda. Proceedings, American Philosophical Society, xlvii: 97-110, with 3 plates, 1908; The Comparative Leaf Structure of the Strand Plants of New Jersey. Proceedings Ameri- can Philosophical Society, xlviii: 72-89, with 3 plates, 1909. t Linnaean Society's Journal — Botany, xxxvii: 333-406, October, 1906. t Liv: 265-305. 259 26o VEGETATION OF THE NEW JERSEY PINE-BARRENS make up the bulk of the leaf in cross-section. Succulent leaves are those in which water storage is accomplished. The depression of the stomata, the development of a thick cuticle, the presence of a hypodermis of thick-walled cells, the presence of hairs and the formation of air-still chambers by a rolling or a folding of the leaf are all structures which assist in the regulation of transpiration. The following is a classification of the different leaf structures of 55 species of pine-barren plants. The species are arranged systematically under each head. Thin Cuticle. — Myrica carolinensis, Quercus ilicifolia, Q. prinoides, Q. stellata, Sassafras variifolium, Azalea viscosa (West Plain), Andro- meda (Neopieris) mariana, Gaylussacia dumosa, Vaccinium vacillans, V. atrococcum, V. macrocarpon. Thick Cuticle. — Chamaecyparis thyoides, Pinus rigida, Xerophyllum asphodeloides, Alnus rugosa, Quercus ilicifolia (East Plain), Q. mary- landica, Corema Conradii, Ilex glabra, Kalmia latifolia, Chamaedaphne (Cassandra) calyculata, Dendrium (Leiophyllum) buxifolium, Epigaea repens, Arctostaphylos uva-ursi, Gaultheria procumbens. Thick-walled Epidermis. — Schizaea pusilla, Pinus rigida, Chamae- cyparis thyoides, Xerophyllum asphodeloides, Pyxidanthera barbulata. Hypodermis Present. — Pinus rigida, Chamaecyparis thyoides, Xero- phyllum asphodeloides, Castalia odorata (in patches), Corema Conradii. Single Row of Palisade. — Pteris aquilina, Chamaecyparis thyoides, Comptonia asplenifolia, Quercus marylandica [Lakehurst], Q. prinoides, Q. stellata, Arenaria caroliniana, Lespedeza frutescens, Hudsonia eri- coides, Acer carolinianum. Sassafras variifolium, Clethra alnifolia [Cedar Swamp], Azalea viscosa, Gaylussacia frondosa, Vaccinium atrococcum, Liatris graminifolia, Eupatorium verbenaefolium. Two or More Rows of Palisade. — Sagittaria longirostra, Smilax glauca, S. laurifolia, Orontium aquaticum, Myrica carolinensis, Alnus rugosa, Quercus ilicifolia, Q. marylandica [West Plain], Q. prinoides [Shamong], Q. stellata [Shamong], Magnolia glauca, Castalia odorata, Tephrosia (Cracca) virginiana, Ilex glabra. Euphorbia ipecacuanhae, Corema Conradii, Arctostaphylos uva-ursi, Azalea viscosa, Chamae- daphne (Cassandra) calyculata, Clethra alnifolia (pine woods), Epigaea repens, Gaultheria procumbens, Gaylussacia dumosa, G. resinosa, Kal- mia latifolia, Dendrium (Leiophyllum) buxifolium, Andromeda (Lyonia) ligustrina, Andromeda (Neopieris) mariana, Vaccinium corymbosum, V. macrocarpon, V. pennsylvanicum, V. vacillans, Chrysopsis falcata, C. mariana, Solidago stricta, Pyxidanthera barbulata, Melampyrum lineare. VEGETATION OF THE NEW JERSEY PINE-BARRENS 261 Stomatal Guard Cells at Surface. — Pteris aquilina, Sagittaria longi- rostra, Comptonia asplenifolia, Alnus rugosa, Quercus ilicifolia, Q. marylandica, Arenaria caroliniana, Magnolia glauca, Sassafras varii- folium, Tephrosia (Cracca) virginiana, Acer carolinianum, Hudsonia ericoides, Azalea viscosa, Clethra alnifolia [Cedar Swamp], Epigaea repens, Gaultheria procumbens, Gaylussacia dumosa, G. frondosa, G. resinosa, Andromeda (Lyonia) ligustrina, Andromeda (Neopieris) mari- ana, Vaccinium atrococcum, V. corymbosum, V. macrocarpon, V. pennsylvanicum, V. vacillans, Pyxidanthera barbulata, Chrysopsis falcata, C. mariana, Eupatorium verbenaefolium, Solidago stricta. Stomatal Guard Cells Projecting. — This refers to the projection of the guard cells beyond the general epidermal surface. Schizaea pusilla, Myrica carolinensis, Corema Conradii, Clethra alnifolia (pine woods) Melampyrum lineare. Stomatal Guard Cells Slightly Depressed. — Smilax glauca, S. laurifolia, Castalia odorata, Lespedeza frutescens, Euphorbia ipecacuanhae, Ilex glabra, Chamaedaphne (Cassandra) calyculata, Dendrium buxifolium, Liatris graminifolia. Stomatal Guard Cells Deeply Depressed. — Pinus rigida, Orontium aquaticum (upper), Xerophyllum asphodeloides, Arctostaphylos uva- ursi, Kalmia latifolia. Leaves Glabrous. — Pteris aquilina, Pinus rigida, Smilax glauca, S. laurifolia, Xerophyllum asphodeloides, Alnus serrulata, Euphorbia ipe- cacuanhae, Arctostaphylos uva-ursi, Clethra alnifolia, Epigaea repens, Gaultheria procumbens, Kalmia latifolia, Andromeda (Neopieris) mari- ana, Vaccinium pennsylvanicum, V. vacillans, Pyxidanthera barbu- lata. Leaf Hairy on Under Side. — Quercus ilicifolia, Q. marylandica, Q. prinoides, Castalia odorata. Magnolia glauca, Lespedeza frutescens. Azalea viscosa [West Plain], Gaylussacia frondosa, Vaccinium atrococ- cum, V. corymbosum. Leaf Hairy on Both Sides. — Comptonia asplenifolia, Myrica carolinen- sis, Quercus stellata, Sassafras variifolium, Corema Conradii, Tephrosia (Cracca) virginiana, Chamaedaphne (Cassandra) calyculata, Gaylus- sacia dumosa, G. resinosa, Andromeda (Lyonia) ligustrina, Melampy- rum lineare, Chrysopsis mariana, Eupatorium verbenaefolium, Liatris graminifolium, Solidago stricta. Epidermal Surface Papillate. — Schizaea pusilla, Orontium aquaticum, Smilax glauca, Alnus rugosa, Comptonia asplenifolia, Lespedeza frutes- cens, Tephrosia (Cracca) virginiana, Azalea viscosa, Chamaedaphne (Cassandra) calyculata, Andromeda (Neopieris) mariana. 262 VEGETATION OF THE NEW JERSEY PINE-BARRENS Leathery Leaf. — Smilax laurifolia, Myrica carolinensis, Quercus ilici- folia, Q. marylandica, Q. stellata, Castalia odorata, Ilex glabra, Arcto- staphylos uva-ursi, Chamaedaphne (Cassandra) calyculata, Epigaea repens, Gaultheria procumbens, Gaylussacia dumosa, G. resinosa, Kal- mia latifolia, Dendrium (Leiophyllum) buxifolium, Vaccinium macro- carpon, V. pennsylvanicum. IViry Leaf. — Schizaea pusilla, Chamaecyparis thyoides, Pinus rigida, Xerophyllum asphodeloides, Arenaria caroliniana, Corema Conradii, Hudsonia ericoides, Pyxidanthera barbulata. Deciduous Leaf. — Pteris aquilina, Smilax glauca, Comptonia aspleni- folia, Myrica carolinensis, Alnus rugosa, Quercus ilicifolia, Q. mary- landica, Q. prinoides, Q. stellata, Castalia odorata. Magnolia glauca, Tephrosia (Cracca) virginiana, Acer carolinianum, Sassafras variifolium, Azalea viscosa, Clethra alnifolia, Gaylussacia dumosa, G. frondosa, G. resinosa, Andromeda (Lyonia) ligustrina, Andromeda (Neopieris) mariana, Vaccinium atrococcum, V. corymbosum, V. pennsylvanicum, V. vacillans. Evergreen Leaf. — Chamaecyparis thyoides, Pinus rigida, Smilax lauri- folia, Xerophyllum asphodeloides. Ilex glabra, Arctostaphylos uva-ursi, Chamaedaphne (Cassandra) calyculata (?), Epigaea repens, Gaultheria procumbens, Kalmia latifolia, Dendrium buxifolium, Vaccinium macro- carpon, Pyxidanthera barbulata. Latex Tubes. — Euphorbia ipecacuanhae. Mucilage. — Quercus marylandica [West Plain], Castalia odorata, Lespedeza frutescens. Sassafras variifolium. Rhomhoidal Crystals {Calcium Oxalate). — Smilax glauca, Lespedeza frutescens, Andromeda (Neopieris) mariana, Vaccinium corymbosum. Sphaerocrystals. — Orontium aquaticum, Quercus ilicifolia, Arenaria caroliniana, Acer carolinianum. Ilex glabra, Chamaedaphne (Cassandra) calyculata, Clethra alnifolia, Epigaea repens, Gaultheria procumbens, Gaylussacia resinosa, Ilex glabra, Kalmia latifolia. Idioblasts. — Castalia odorata. Resin Hairs. — Myrica carolinensis, Gaylussacia resinosa. Resin Canals. — Pinus rigida. The following classificatory heads are based on the general structure of the leaf, i. e.,, the arrangement of the palisade and the spongy paren- chyma. Diphotophyll. — Pteris aquilina, Orontium aquaticum, Sagittaria longi- rostra, Smilax glauca, S. laurifolia, Comptonia asplenifolia, Myrica carolinensis, Alnus rugosa, Quercus ilicifolia, Q. marylandica, Q. pri- noides, Q. stellata, Castalia odorata. Magnolia glauca. Sassafras varii- folium, Lespedeza frutescens, Tephrosia (Cracca) virginiana, Corema VEGETATION OF THE NEW JERSEY PINE-BARRENS 263 conradii, Acer carolinianum, Ilex glabra, Hudsonia ericoides, Arcto- staphylos uva-ursi, Azalea viscosa, Chamaedaphne (Cassandra) caly- culata, Epigaea repens, Gaultheria procumbens, Gaylussacia dumosa, G. frondosa, G. resinosa, Dendrium buxifolium, Andromeda (Lyonia) ligustrina, Andromeda (Neopieris) mariana, Vaccinium atrococcum, V. corymbosum, V. macrocarpon, V. pennsylvanicum, V. vacillans, Pyxidanthera barbulata, Melampyrum lineare, Chrysopsis falcata, C. mariana, Eupatorium verbenaefolium, Liatris graminifolia. Diplophyll. — Chamaecyparis thyoides, Arenaria caroliniana, Euphor- bia ipecacuanhae, Solidago stricta. 5/aMro^/;>'//.— Smilax glauca (?), S. laurifolia (?), Arctostaphylos uva- ursi (incipient), Kalmia angustifolia, K. latifolia. Spo7igophyIl. — Pyxidanthera barbulata (?). Sclerophyll. — Schizaea pusilla, Pinus echinata, P. rigida, Xerophyllum asphodeloides. Roll Leaf Structure. — Corema Conradii. Insectivorous Leaf. — Drosera rotundifolia. Detailed Microscopic Structure of Leaves The sections of leaves and other parts, which are figured and described in this chapter, were made free hand with a razor, then stained with Bismarck brown and mounted for permanency in Canada balsam. The drawings were made by Edinger's drawing and projection apparatus, manufactured by E. Leitz, Wetzlar. A description of this instrument will be found in the trade circular of that firm and need not be re- counted here. The drawings were first made in lead pencil and finished in India ink. Using the same magnification for each drawing, the sections are drawn to the same scale, a highly desirable end in making a comparison of leaf section with leaf section. In order to attain the 'greatest accuracy, each drawing before it was inked was compared with the corresponding section under the microscope, and the following de- scriptions were written with the drawings and the microscopic sections before me. The drawings of stomata were not m.ade to scale. The source of the material is given in each instance where such data have been kept.* Pteris aquilina [Lakehurst]. — The bracken-fern has dull-green, ter- nately compound fronds at the summit of an erect, stout petiole (2 to 9 dm. high) from a creeping root-stock. Both sides of the leaf are gla- brous with thin-walled epidermal cells of unequal size. The guard cells * In the reproduction for printing, each drawing has been reduced one-half. 264 VEGETATION OF THE NEW JERSEY PINE-BARRENS Figure 206 Pteris aquilina, Lake- hurst. of the stomata on the lower side project sHghtly. There is a single row of palisade cells. A typic diphotophyll (Fig. 206). Schiiaea pusilla [Cedar Swamp, Shamong]. — The curly-grass fern is a small inconspicuous plant with distinct sterile and fertile fronds that are very slen- der, linear and wiry, usually curled. As shown by the figure, the structure of the stipe of both fronds is essentially alike, that of the fertile being larger and thicker than that of the sterile. The epider- mal cells have thick outer walls that are distinctly papillate, with a slight boss, or projection, on each epidermal cell. The stomata are confined to the lower or curved surface, and the guard cells project considerably beyond the general surface. The chlorenchyma cells have slightly thicker walls than usual, and in the leaf-trace bundle the phloem occurs on the abaxial side of the bundle, which is usually roughly elliptic in cross-section with its long axis tangential to the stele.* A spongophyll (Fig. 207). Chamaecyparis thyoides. — The white-cedar has small, scale-like or awl-shaped, closely appressed, imbricated leaves often with a small gland on the back. The epidermis is protected by a thick cuticle, and is supported by a single row of hypo- dermal sclerenchyma cells. A single row of pali- sade cells is found on both sides of the leaf. The spongy parenchyma is rather compact. A diplophyll (Fig. 208). Pinus rigida [Lakehurst]. — The leaves of the pitch-pine are in fascicles of three. Their length varies from 5 to 12 centi- meters, and they are dark-green in color. The thick-walled epidermal cells are cov- ered with a cuticle, and they are rein- forced by silvery, thick-walled hypoder- mal sclerenchyma, which is represented by two or three rows of cells that increase in number at the margin of the leaf. The guard cells of the stomata are thick-walled and are depressed below the Figure 207 Schizaea pusilla, Cedar Swamp, Shamong, June 28, 19 10. Figure 208 Chamaecyparis thyoides, Cedar Swamp, Shamong (Chatsworth), June 28, 1910. * SiNNOTT, Edmund W.: The Evolution of the Filicean Leaf-trace. Annals of Botany, xxv: 171, January, 191 1. VEGETATION OF THE NEW JERSEY PINE-BARRENS 265 surface, until they are found on a level with the first row of hypodermal sclerenchyma cells. The walls of the rounded, mesophyll parenchyma cells are ruminate by projections of the wall into the cell cavity. Three Figure 209 Pinus rigida, Lakehurst. resin canals are found and there are two leaf-trace bundles surrounded by large parenchyma cells and a bundle sheath. A leaf from a tree found on the Upper Plain is thicker transversely, the parenchyma cells are larger, and the resin canals larger and more open (Fig. 209). Orontium aqiiaticum [Shamong]. — The golden-club has long-petioled, entire, ob- long, and nerved leaves. The thin-walled epidermal cells of the petiole are vertically placed. The parenchyma cells are spaced widely by large, rounded, intercellular spaces, so that a reticulum, or net, of these cells is formed with the collateral vascular bundles attached here and there to several meshes of this parenchymatous net. A transverse section of the leaf-blade shows the upper epidermal cells strongly papillate and interspersed with stomata, the guard cells of which are deeply depressed. There are at least two rows of palisade cells, and the spongy parenchyma has large, irregular, intercellular spaces. Sphaerocrystals are present in these cells. The lower epidermal cells are large, with their outer walls smooth and not papillate. A diphotophyll (Figs. 210 and 211). Figure 2 10 Orontium aquaticum. Swamp, Shamong, N. 28, 19 10. Cedar . June 266 VEGETATION OF THE NEW JERSEY PINE-BARRENS Sagiiiaria longirostra [Shamong]. — This species of arrow-leaf has broadly ovate-oblong, obtusish, sagittate leaves. The epidermal cells of both leaf surfaces are large and thin-walled. Stomata level with the surface are present on both leaf surfaces and opposite large intercellular spaces. There are 3 to 4 rows of palisade. A diphotophyll (Fig. 212). Smilax glauca. — The saw-brier is a climbing vine with terete branches and somewhat four- angled, glaucous branchlets armed with scattered prickles. The leaves Figure 212 Sagittaria longirostra, Shamong, August 29, 1910. are ovate and glaucous beneath. The upper epidermis consists of thin-walled, brick-like cells. The Figure 2 1 1 Petiole of Orontium aquaticum, Cedar Swamp, Shamong (Chatsworth), June 28,1910. Figure 213 Smilax glauca, East Plain, August 31, 1 9 10. epidermal cells of the lower side are papillate and the guard cells of the stomata on the lower surface are depressed only slightly. There are two rows of short, rounded palisade cells and the short, transversely placed, spongy parenchyma cells are aligned with those of the palisade. Rhomboidal crystals are present. A diphotophyll simulating a stauro- phyll (Fig. 213). VEGETATION OF THE NEW JERSEY PINE-BARRENS 267 Figure 214 Smilax laurifolia, Cedar Swamp, Shamong, June 28, 19 10. Smilax laurifolia [Cedar Swamp, Shamong]. — This woody, cHmb- ing plant has thick, evergreen, coriaceous leaves (6-12 cm. long). The regular, brick-like, upper and lower epidermal cells are covered by a thick cuticle. The stomata have their guard cells depressed slightly below the surface. There are two to three rows of palisade cells and the loose parenchyma cells are aligned with the palisade. A diphotophyll, or a potential staurophyll (Fig. 214).* Xerophyllum asphodeloides. — The turkey's- beard is an herb with a short stem from a thick, tuberous root-stock bearing a crown of long, needle-shaped, wiry leaves. The upper, flat sur- face of the leaf has small, thick-walled epidermal cells, reinforced by three to several rows of hypodermal sclerenchyma connected with each of the several bundles of the leaf. The lower curved surface of the leaf is provided with a few large, stiff, tooth-like hairs. The epidermal cells are thick-walled, supported by several rows of hypodermal sclerenchyma. The stomata, found on both surfaces, have thick-walled, depressed guard cells covered by two projecting tooth-like epidermal cells. The bundles, which are of the usual monocotyledonous, collateral type, are surrounded completely with a bundle-sheath of sclerenchyma. The mesophyll parenchyma cells, which are hex- agonal or pentagonal in shape, are very com- pact and the walls are thicker than the walls of ordinary fundamen- tal tissue. A sclerophyll (Fig. 215). Compionia asplenifo- lia [Lakehurst]. — The sweet-fern is a shrub 3 to 6 decimeters tall, with sweet-scented, fern-like, lanceolate pinnatifid leaves that are deciduous. Both surfaces of the leaves are hairy, with straight, rather stiff, unicellular hairs. The epidermal cells are slightly * See Kearney, T. H.: Contributions U. S. National Herbarium, 5: 486, f 86, 1901. Figure 2 15 Xerophyllum asphodeloides, between Shamong and the Plains, June 28, 1910. 268 VEGETATION OF THE NEW JERSEY PINE-BARRENS Figure 216 Comptonia aspleni- folia, Cedar Swamp, Shamong, June 28, 19 10. Figure 217 Comptonia asplenifolia, West Plains, June 28, 19 10. papillate and the guard cells on the lower side are at the surface. There is a single row of pali- sade in the leaves from Lake- hurst and two rows in the leaves from the Upper Plain, thus increasing the thickness of the leaves. In both leaves there is open loose paren- chyma. Adiphotophyll (Figs. 216 and 217). Myrica carolinensis [Cedar Swamp, Shamong]. — The bay- berry is a shrub 1-2 meters tall, with oblong, entire, or crenately toothed leaves. The upper epi- dermal cells are thin-walled, with a cuticle and an occasional long, stiff unicellular hair. There are 2 to 3 rows of. palisade cells and the spongy layer is a net of parenchyma cells. The lower epidermis has similar hairs and stomata, the guard cells of which project slightly. Occasional multicellular hairs occur in depressions of the lower epidermis. A diphotophyll (Fig. 218). Alnus riigosa { = A. serrulata) [Shamong]. — The smooth alder is a shrub or small tree with ob- ovate, sharply serrate leaves, smooth or slightly pubescent beneath. The regular, brick-like, up- per epidermal cells are covered with a thin cuticle. There are 3 rows of palisade cells. The spongy parenchyma is open. The guard cells of the sto- mata on the lower side are at the surface. A diphotophyll (Fig. 219). Quercus ilicifoUa{^Q. nana) [Lakehurst, Shamong, Lower Plain].— The bear or scrub oak is a shrub with obovate leaves, wedge-shaped at the base, angularly 3 to 7 lobed, white, downy beneath and glabrous above. A thin cuticle is present on the upper sur- face of the leaf of the plants from Shamong and the Lower Plain. The epidermal cells are thin-walled, and the bent, unicellular hairs of the lower surface are clustered with four or five hairs arising from a single point of attachment with occasional multicel- The under surface of the Shamong leaf is more Figure 218 Myrica carolinensis, Cedar Swamp, Sha- mong, June 28, 1910. Figure 219 Alnus rugosa. Cedar Swamp, Shamong, June 28, 1910. lular capitate hairs. VEGETATION OF THE NEW JERSEY PINE-BARRENS 269 Figure 220 Quercus ilicifolia. East Plain, August 31, 1910. hairy than in the Lakehurst leaf. There are two rows of paHsade cells in the leaf of the Lakehurst plant and three rows in the leaf of the Shamong plant. The parenchyma is rather compact. The stomata have their guard cells at the surface. Sphaero- crystals are present in some of the cells. A dipho- tophyll (Fig. 220). Quercus marylandica [Lakehurst and Upper Plain]. — The black-jack oak is a small tree with broadly wedge-shaped leaves, widely dilated and somewhat 3 to 5 lobed at the summit, rusty pub- escent beneath, shining above. The large cells of the upper epidermis are protected by a thick cuticle, and in the epidermal cells from leaves gathered on the Upper Plain there is found a gum-resin-like substance absent in the cells of the leaves from Lakehurst. There is a single row of true palisade cells in the Lakehurst leaves and two rows in the leaves of the specimen from the Upper Plain, and the lower epidermis displays the stomatal guard cells at the surface. The large, unicellular hairs are present on the lower surface and also branched, stalked, and capitate multicellular hairs. The spongy parenchyma is rather open. A diphotophyll (Fig. 221). Quercus prinoides [Lakehurst and Shamong]. — This species of oak is a low, spreading bush with undulate leaves covered beneath by a close white tomentum. The upper epidermal cells, which are thin-walled, are pro- tected by a clearly defined cuticle. There is a single row of palisade, while in the leaf from the pine-barrens at Shamong there are two rows and the spongy parenchyma is rather compact. The stomata on the under side have their guard cells at the surface. The hairs characteristic of the under side are straight, tufted and unicellular. A diphoto- phyll (Fig. 222). Quercus stellata ( = Q. minor) [Lakehurst, Shamong, Lower Plain].— The post-oak is a tree with thick, obovate, lyrate, pinnatifid leaves, sinuately cut into 5 to 7 rounded lobes, the upper of which are much larger and broader. The upper surface is rough, with hairs stellately arranged, and the lower surface is brownish-downy with curved stiff hairs in tufts. A cuticle is present. A single row of palisade is found in the leaves from Lake- FlGURE 221 Quercus marylandica, Lakeiiurst. C^3) Figure 222 Quercus prinoides, Sha- mong, June 28, 191 1. 270 VEGETATION OF THE NEW JERSEY PINE-BARRENS Quercus Figure 223 stellata, Shamong, June 28, 191 1. hurst and Lower Plain and three in the leaf from Shamong. The guard cells of the stomata on the lower side are flush with the surface. A diphotophyll (Fig. 223). Quercus veliitina [Lakehurst]. — The quercitron-oak is a tree with leaves variously divided. Both surfaces of the leaf are smooth, but the pubes- cence is in tufts in the axils of the veins beneath. The upper epidermal, thin-walled cells are provided with a cuticle. The palisade cells are found in a single row, and the stomata have guard cells flush with the surface. A diphotophyll. Arenaria caroliniana [Shamong]. — The pine-barren sandwort is a densely tufted plant with closely imbricated but spreading, awl-shaped leaves. The thin-walled epider- mis, which covers the surface of the approx- imately semicircular leaf in cross-section, is without a cuticle. Palisade cells are devel- oped on both surfaces and the rather com- pact loose parenchyma with sphaerocrystals surrounds a centrally placed bundle system with strongly marked bundle sheath. Two small patches of sclerenchyma occupy the corners of the leaf. The guard cells of the stomata are at the surface on both sides of the leaf. A diplophyll (Fig. 224.). Casialia (Nymphaea) odorata. — The leaves of the water-lily are orbic- ular, 0.5 to 2.2 dm. in diameter, deeply cordate at the base, with the margin entire. The under surface is a deep crimson-brown. The microscopic structure of the petiole is as follows: The epidermis consists of thin- walled parenchyma cells, the outer wall rounded and projecting. Thin-walled hairs are found scattered over the surface. Be- neath the epidermis the hypodermal cortex cells are slightly collenchymatous, and thin- walled parenchyma cells are interspersed between large, open intercellular canals into some of which star-like idioblasts project, their walls being roughened with crystalline deposits. The bundles are collateral with the phloem external and the xylem inter- nal. A round canal accompanies invariably each bundle. Mucilage cells are present also. Figure 224 Arenaria caroliniana, Shamong, June 28, 191 1. VEGETATION OF THE NEW JERSEY PINE-BARRENS 271 Figure 225 Castalia (Nymphaea) odorata, Cedar Swamp, Sha- mong, June 28, 1910. The microscopic structure of the leaf blade is of interest. The upper epidermis of thin-walled cells alone shows the stomata with their guard cells depressed slightly be- low the surface. There are four to five rows of palisade cells and a wide area of spongy parenchy- ma with extremely large intercellular air chambers, crossed by chains, or bridges, of parenchyma cells. Into these air spaces project large, thick-walled, star-shaped id io blasts, which penetrate also to the upper epidermal cells between those constitut- ing the palisade. The col- lateral bundles with xylem uppermost are tied to the lower epidermis by patches of sclerenchyma. Peculiar flattened epi- dermal hairs are found on the under surface. A diphotophyll (Figs. 225 and 226). Magnolia glauca { = M. virginiana) [ S h a m o n g and Lower Plain]. — The sweet-bay is a tree with broadly lanceolate to elliptic obtuse leaves, 8 to 1 5 cm. long, glaucous beneath. The small up- per and lower epider- mal cells are without cuticle. Long, straight, 3- to 4-celled, silky hairs are found on the lower side. The guard cells There are two rows of palisade cells and Figure 226 Castalia (Nymphaea) odorata, Cedar Swamp, Shamong, June 28, 1910. are level with the surface. 272 VEGETATION OF THE NEW JERSEY PINE-BARRENS Figure 227 Magnolia glauca, Cedar Swamp, Sha- mong, June28, 19 10. Figure 228 Sassafras officinale, Lakehurst. the spongy parenchyma is somewhat compact. A diphotophyll (Fig. 227). Sassafras variifolium [Lakehurst]. — The sassafras is a low tree in the pine-barrens with ovate, entire, i- to 3-lobed, pubes- cent, mucilaginous leaves. The upper epidermis, which is cuticularized, is provided sparingly with short, stiff, unicellular hairs. There is a single row of open palisade tissue with large, rounded, lysigenous cavities* filled with mucilage. The loose parenchyma cells are fiat and irregular in shape. The lower epidermis of thin-walled cells shows a number of straight, or slightly bent, unicellular hairs. The guard cells of the stomata are at the surface of the under side. A diphotophyll (Fig. 228). Drosera rotundifolia. — The sundew is a plant of such unusual biologic interest that an enlarged drawing of one of its leaf tentacles is added. The stalk of the tentacle is supplied with a few spiral tracheae. A plexus of spiral tracheids in the enlarged terminal portion of the tentacle is characteristic. This plexus is covered with modi- fied epidermal and subepidermal cells which secrete a glairy fluid to which small insects adhere. When stimulated by insects, the tentacles curve inwardly over the central part of the leaf, thus trapping the insect, the soft parts of which are absorbed gradu- ally by the digestive action of the secretions that are formed for that purpose (Fig. 229). Lespede^a frutescens [Shamong]. — The stems of this plant (1.5 to 7 dm. tall) have compound trifoli- ate leaves with oval to oblong firm leaflets with finely appressed pubescence. The epidermal cells are thin-walled, and on the under surface some of them are papillate. There is a single row of long palisade cells. Rhomboidal crystals of calcium oxa- late are present in some of the cells. Some of the spongy parenchyma cells apparently have a mucilaginous material which stains deeply with Bismarck brown. The guard cells are depressed * In Solereder's Systematic Anatomy of Dicotyledons, 1 1 : 703 (190S), these are referred to in the order as "enlarged sac-like cells." Figure 229 Tentacle of Drosera ro tundifolia. VEGETATION OF THE NEW JERSEY PINE-BARRENS 273 Figure 230 Lespedeza frutes- cens, Shamong, Au- gust 29, 1910. slightly. Straight hairs occur on the under leaf surface. A diphoto- phyll (Fig. 230). Tephrosia (Cracca) virginiana [Lakehurst, Upper Plain and Lower Plain]. — -The goat's-rue is a per- ennial plant with an erect stem, 3 to 6 dm. high, covered with silky villous hairs when young. The yellowish-white flowers, marked with purple, are in dense racemes. The leaves are compound, and there are 17 to 19 linear-oblong leaflets. The upper epi- dermis consists of thin-walled, arched cells raised as papillae in some places, in others replaced by the stiff and slightly curved, unicellular hairs. There are 2 rows of palisade cells in the leaves from plants gathered on the Upper Plain and the pine-barrens at Lakehurst, and three in the leaves from a plant growing on the Lower Plain. The loose parenchyma consists of rounded cells with large intercellular spaces. The lower epidermal cells are thin-walled, and the guard cells of the stomata on both the upper and lower sides are flush with the surface. A typic diphoto- phyll (Fig. 231). Euphorbia ipecacuanhae [Shamong and Lower Plain]. — This spurge has leaves that vary extremely in form and color. The shape varies from obovate, or oblong, to narrowly linear and glabrous. The color varies from green to deep-red. The epidermal cells on both the upper and lower sides of the leaf are thin-walled. The guard cells are depressed on both surfaces, and a single row of palisade cells is found on the upper and lower sides, the lower being more prominent in the leaf from Shamong, with spongy parenchyma between. Latex tubes are found in all cross-sections. A diplophyll (Fig. 232). Corema Conradii [Lower Plain]. — The broom-crow- berry has nearly verticillate, linear, heath-like leaves. In cross-section these leaves show a typic roll struc- ture, with both margins rolled together until only a narrow, hair-protected slit is left. The whole ex- posed outer surface is covered with a thick cuticle, beyond which project multicellular capitate hairs. The chambered inner surface displays two kinds of hairs — multicellular, capitate hairs and straight, unicellular hairs, which are found most numerously along the two inturned edges of the leaf, thus guarding Figure 23 i Tephrosia virginiana, Lakehurst. Figure 232 Euphorbia ipecacu- anhae, East Plain, August 3 I, 1910. 274 VEGETATION OF THE NEW JERSEY PINE-BARRENS Figure 233 Corema Conradii, East Plain, August 20, 191 1. the slit-like opening between the leaf margins. The stomata are found on the lower, or inner, surface of the leaf and the guard cells project beyond the general surface. Be- neath the upper epidermal cells are large, looped cells filled with a ma- terial suggesting a gum-resin. This material stains deeply with Bis- marck brown. The palisade cells arearranged in 2 to 3 layers and the spongy parenchyma is open with large, irregular, intercellular spaces. A diphotophyll (Fig. 233). Ilex glabra [Shamong, between Shamong and the Plains, Lower Plain]. — The inkberry is a shrub 6 to 9 decimeters tall, with oblong, smooth leaves. A study of sections of leaves gathered from plants in the above localities enables one to note the influence of environment on leaf structure. The upper and lower epider- mal cells of leaves from all three localities are protected by a thick cuticle. The stomata, confined to the lower surface, have their guard cells depressed the thickness of the cuticle. Sphaerocrystals are present in the spongy parenchyma of the leaves from the three localities. Figure 234 Ilex glabra between Shamong and the Plains, June 28, 1910. Figure 235 Ilex glabra, Cedar Swamp, Shamong, June 28, 1910. Figure 236 Acer caroiinianum, Cedar Swamp, June 28, 1910. arched slightly. The leaf from the white-cedar swamp at Shamong displays three rows of palisade cells and a very open spongy parenchyma. This leaf is thinner than the other two. The leaves from the Lower Plain and from east of Shamong have four rows of palisade cells and a more compact loose parenchyma. A diphotophyll (Figs. 234 and 235). Acer caroiinianum [Cedar swamps, Shamong and edge of Upper Plain]. — The Carolinian red maple has small, obovate leaves, with three short lobes. The upper epidermal cells are large, with the upper wall The lower epidermal cells are thin-walled and papillate. VEGETATION OF THE NEW JERSEY PINE-BARRENS 275 The stomata, confined to the lower surface, have superficial, obliquely- placed guard cells. There is a single row of palisade cells and the spongy parenchyma is semi-compact. A diphotophyll (Fig. 236). Hudsonia ericotdes [Upper Plain]. — This plant is a bushy, heath-like, small shrub, with slender, awl-shaped leaves which vary in outline in different parts of the serial cross-sections. The epidermal cells are thin- walled, with the stomata at the surface. There is a single row of pali- sade cells and in front of each stoma there is an intercellular chamber developed in the otherwise rather compact, spongy parenchyma. The bundle is placed centrally. A few straight hairs are scattered over the surface. The leaf is a diphotophyll. The stem section is circular, and the epidermis of thin-walled cells develops numerous long, silky, uni- cellular hairs interspersed with multicellular, stalked, capitate hairs. The bundle system is central and is surrounded by a cortex of thin-walled parenchyma cells (Figs. 237 and 238). Figure 237 Stem of Hudsonia ericoides, West Plain, June 28 19 10. Figure 238 Leaf of Hudsonia ericoides, West Plain, June 28, 1910. Ardostaphylos uva-ursi [Shamong, Upper and Lower Plains]. — The bearberry is a trailing plant with thick, evergreen, obovate or spat- ulate, entire, smooth leaves. Both the upper and the lower epidermal cells are covered with a thick, yellowish cuticle with stomata on the lower surface only. The large, thick-walled guard cells are depressed an amount equal to the thickness of the cuticle. There are five to six rows of palisade cells and the loose parenchyma is open with many of the cells directed vertically, so that the leaf becomes incipiently a staurophyll. The leaves of the pine-barren plant collected at Shamong are thinner by one row of palisade cells than are those leaves gathered in the Upper and Lower Plains (Fig. 239). The leaves of the bearberry contain a glucoside, arbutin. It has been 276 VEGETATION OF THE NEW JERSEY PINE-BARRENS Figure 239 Arctostaphylos uva-ursi, West Plain, June 28, 1 9 10. discovered by a new method of microscopic analysis, known as the microsubHmation method, that when the powdered leaves of Arcto- staphylos are heated in a watch-glass and the sublimable principles collected on microscopic slides, the arbutin is split up into hydroqui- none, which is deposited on the slides in several crystalline forms, character- istic of the plant. Weevers considers that arbutin, as a glucoside, is a reserve food and is stored in the leaves, and when the new leaves are formed in the spring, it is split by a suitable enzyme into sugar and hydroquinone. The sugar is used up, and the hydroquinone remains behind and combines with more sugar, so that the autumn leaves once more contain arbutin.* By this method arbutin (hydroquinone gluco- side) has been detected in Epigaea repens, Gaultheria procumbens and Kalmia angustifolia, a description of which plants follows. Aialea {Rhododendron) viscosa [Cedar swamps, Shamong, between Shamong and Upper Plain, edge of Upper Plain]. — The white swamp- honeysuckle is a shrub with oblong-obovate, smooth leaves, bristly along the margins and the midrib. The leaf from the white-cedar swamp at Shamong displays large thin-walled upper epidermal cells, a single row of palisade and a lower epidermis, with the guard cells of the stomata at the surface. The stomata are found only on the lower side. In the leaf from a plant collected between Shamong and the Lower Plain the thickness of the leaf has in- creased. A few of the upper epidermal cells are papillate, the palisade cells are longer, and the spongy parenchyma shows a larger number of layers and is more compact. In the leaf of a plant which grew at the edge of a white-cedar swamp along the Upper Plain the number of palisade layers is three and the spongy parenchyma is open. Glandular capitate hairs are found sparingly on the lower epidermis. A diphotophyll (Fig. 240). * MoLiscH, Hans: Mikrochemie der Pflanze, 1913: 165-166; Haas, Paul, and Hill, T. G.: An Introduction to the Chemistry of Plant Products, 1913: 171; Tunmann, O.: Ueber Folia uvae-ursi und den mikrochem. Nachweis des Arbutins. Pharm. Zentralb., 1906, XLvii, No. 46; Tunmann, O.: Pflanzen Mikrochemie, 1913: 355. Figure 240 Azalea viscosa, West Plain, edge of Cedar Swamp, August 3 I, 1910. VEGETATION OF THE NEW JERSEY PINE-BARRENS 277 Figure 241 Cassandra calycu- lata, Cedar Swamp, Shamong, June 28, 1910. Chamaedaphne (Cassandra) calyculata. — The leather-leaf is a low shrub with coriaceous evergreen leaves. A thick cuticle covers the small, thin- walled upper and lower epidermal cells. There are two to four rows of palisade cells. The spongy par- enchyma is rather compact, with occasional sphaero- crystals. The stomata, found on the lower side of the leaf, have their guard cells depressed slightly and protected by tooth-like projections of the adjoining epidermal cells. Saucer-shaped, peltate hairs give to the lower leaf surface a rusty, scurfy appearance. Such hairs are distributed sparingly over the upper surface. A diphotophyll (Fig. 241). Clethra alnifoUa [White-cedar Swamp, Shamong, pine-woods between Shamong and the Plains]. — The sweet pepper-bush is a shrub i to 3 meters tall. The leaves are wedge-obovate, sharply serrate, and prominently veined. The leaf of the white-cedar swamp plant examined has large, rounded, thin-walled upper epidermal cells without stomata and large lower epidermal cells with a single row of palisade cells and open loose parenchyma. The guard cells are at the surface. Sphaerocrystals are present in some of the palisade cells. The leaf of the pine-barren plant has the same large, thin- walled upper and lower epidermal cells, but the guard cells of the stomata on the lower surface project slightly. There are two rows of palisade cells and the spongy parenchyma cells are more compact. Sphaerocrystals are pres- ent. A diphotophyll (Fig. 242). Epigaea repens [Upper Plain and Lower Plain]. — The trail- ing-arbutus is a prostrate plant with evergreen, elliptic, leath- ery leaves on short petioles. The upper epidermis of large cells is protected by a thick yellowish cuticle pierced by scattered stomata, the guard cells of which are flush with the surface. The lower epi- dermal cells are without a cuticle. If we contrast the leaves from the pine-barren plants growing Figure 242 Clethra ainifolia, be- tween Shamong and the Plains, June 28, 1910. cn>cxi3aoooo Figure 243 Epigaea repens (Sun), East Plain, August 31, 1910. Figure 244 Epigaea repens (Shade), Pine-barrens, March 25, 1910. 278 VEGETATION OF THE NEW JERSEY PINE-BARRENS Figure 243 Gaultheria procumbens, East Plain, August 31, 19 10. in the shade and in the sun, we find the shade leaf thinner, with more open spongy parenchyma, while the sun leaf has a cuticle and more compact spongy parenchyma. There are four rows of palisade cells in the leaves from the Lower and the Upper Plains, and the loose parenchyma consists of large, rounded cells. Sphaerocrystals are present. Adiphotophyll (Figs. 243 and 244). Gaultheria procumbens [Lakehurst]. — The tea- berry is a perennial herb with leathery, evergreen, obovate, or oval leaves. The cuticle is thick on the upper surface, and the upper epidermal cells are thin-walled. There are three to four rows of palisade cells and some of the rounded loose paren- chyma cells have sphaerocrystals. The stomata, which are confined to the lower surface, have their guard cells flush with the thin cuticle of that side. The glucoside arbutin is present (see ante). A diphotophyll (Fig. 245). Gaylussacia dumosa [Lakehurst]. — The dwarf huckleberry is a low, bushy perennial, 2 to 15 decimeters tall. Its tardily deciduous leaves are obovate-oblong, thick and shining when old, hairy on both sides. The epidermal cells are thin-walled, the upper with a thin cuticle. There are two rows of palisade cells, the cells of the lower row being slightly divergent. The stomata, as far as the sections available for study show, are confined to the lower side of the leaf and the guard cells are level with the surface. The hairs on both sides are stiff, unicellular, and bent slightly. A typic diphotophyll (Fig. 246). Gaylussacia frondosa [ Lake- hurst, Upper and Lower Plains]. — The dangleberry is a bush with obovate-oblong leaves, finely pu- bescent and glaucous beneath. The upper and lower epidermal cells are large, the lower being arched slightly. The hairs on the under surface are straight and several-celled. There is a single row of palisade cells in the leaves from Lakehurst and the Upper Plain, Figure 246 Gaylussacia dumosa, Lakehurst. Figure 247 Gaylussacia frondosa, West Plain, edge of Cedar Swamp, August 31, 1 9 1 o. Figure 248 Gaylussacia fron- dosa. East Plain, August 31, 19 10. VEGETATION OF THE NEW JERSEY PINE-BARRENS 279 Figure 249 Gaylussacia resinosa, Shamong, June 28, 19 10. and two rows in the leaf of the plant growing on the Lower Plain. The spongy parenchyma consists of rounded cells separated by wide inter- cellular spaces. A diphotophyll (Figs. 247 and 248). Gaylussacia resinosa (= G. haccata) [Shamong, Lower Plain, between Shamong and the Upper Plain]. — The black huckleberry is a much-branched bush, 0.3 to I meter tall, with oval, oblong-ovate, or oblong leaves, thickly covered at first with clammy, resinous globules. The upper and lower epidermal cells are large and thin-walled, bearing in depressions multicellular capitate hairs that are resiniferous on both the upper and lower sides of the leaf. The number of palisade layers varies from i to 2, perhaps 3, and the stomata, confined to the lower surface of the leaf, have their guard cells flush with the surface. Sphae- rocrystals are present in leaves from some localities. A diphotophyll (Fig. 249). Kalmia ayigustifolia [Shamong, Lower Plain]. — The sheep-laurel is an evergreen shrub, i meter high, with opposite to verticillate (in threes), narrowly oblong, obtuse leaves, glabrate underneath. The cells of the lower and of the upper epidermis are protected by a thin cuticle. The upper epidermal cells are larger and more arched than the lower, and the upper wall is thickened considerably. The stomata are confined to the lower surface and the guard cells are depressed only slightly below the surface. As the leaf is a typic stauro- phyll, there are 4 to 5 rows of palisade cells in the leaf from Shamong and 6 rows in the leaf from a twig gathered on the Lower Plain. Sphaerocrys- tals are abundant in the palisade cells. Long, erect, unicellular hairs are present on the upper side of the leaf from the Lower Plain (Fig. 250). Kalmia latifolia [West Plain]. — The laurel has alternate, evergreen, leathery leaves. The cells of the upper and lower epidermis are protected by a thick cuticle. The stomata, confined to the lower surface of the leaf, have their thick-walled guard cells as thick as the cuticle. There are 3 to 4 well-defined rows of palisade cells, and the cells of the spongy Figure 250 Kalmia angustifolia, East Plain, August 31, 1910. Figure 25 1 Kalmia latifolia (Sun), Pine-barrens, March 25, 1910. 28o VEGETATION OF THE NEW JERSEY PINE-BARRENS Figure 252 Kalmia 1 a t i f ol i a , East Plain, August 31, 1910. parenchyma are aligned with those of the paHsade. Sphaerocrystals are present. Sections made from leaves of plants from the pine-barrens (sun) and from the plains agree practically in all details, except that the spongy parenchyma of the pine-barren leaf is meager. A staurophyll (Figs. 251 and 252). Dendriiim {Leiophyllum) biixifolium [Lower Plain]. — The sand-myrtle is a shrub, i to 9 decimeters tall, with oval, or oblong, glabrous, leathery leaves. The upper and lower epidermal cells are protected by a thick cuticle. The guard cells of the stomata, found only on the lower leaf-side, are depressed to the depth of the protective cuticle. There are 3 to 4 rows of palisade cells and the spongy parenchyma cells are separated by large, intercellu- lar spaces. A diphotophyll (Fig. 253). Andromeda {Lyonia, Xolisma) ligustrina [Sha- mong and between Shamong and the Upper Plain]. — The male-berry is a bush, 0.5 to 3 meters tall, with obovate to lanceolate, oblong, serrulate, or entire leaves. The upper and lower epidermal cells, provided with multicellular hairs, are thin- walled. The stomata, found only on the lower leaf-side, have their guard cells at the surface. There are a number of rows of palisade and a rather compact, spongy par enchyma of irregular cells. 254)- Andromeda {Lyonia, Neopieris) mariana. — The stagger-bush is a deciduous shrub with oblong to oval leaves, 3.5 to 8 centimeters long. The upper epidermis of thin-walled cells is protected by a thin cuticle. There are two rows of palisade cells. Rhomboidal crystals of calcium oxalate are present. The lower epidermal cells are slightly papillate and the guard cells of the stomata are at the surface. A diphotophyll. Vaccinium atrococcum [Lower Plain]. — The high black blueberry is a shrub, I to 4 meters tall, with entire leaves, downy or woolly under- neath, even when old. At flowering time the leaves are hardly unfolded. Figure 253 Dendrium (Leiophyl- lum) buxifolium, East Plain, August 31, 1910. A diphotophyll (Fig. Figure 254 Andromeda (Lyonia) ligustrina, between Shamong and the Plains, June 28, 19 10. VEGETATION OF THE NEW JERSEY PINE-BARRENS 281 Figure 253 Vaccinium atrococ- cum. East Plain, August 31, 1910. The upper epidermal cells are protected by a cuticle. The lower epi- dermal cells are smaller in size and some are modified into two kinds of hairs, viz., long, straight, unicellular hairs and capi- tate ones. The stomata are confined to the lower surface and have their guard cells on the epidermal level. There is a single row of palisade cells. A diphotophyll (Fig. 255). Vaccinium corymbosiini [Edge of White-cedar Swamp, Upper Plain].— The high swamp-blueberry is a shrub 1 to 4 meters tall. The leaves are half grown at flowering time, which is later than the preceding species. The ovate to elliptic, lanceolate, smooth leaves are slightly pubescent beneath. The epider- mal cells of the upper and lower sides of the leaf are thin-walled. Stomata with their guard cells level with the surface are found only on the lower side. There are 2 rows of palisade cells and a very open spongy parenchyma. A diphotophyll (Fig. 256). Vaccinium macrocarpon. — The cranberry plant is a trailing one, with oblong-ellip- tic, blunt, evergreen, smooth, to 17 millimeters long, 2 to 8 The large upper and lower epi- There Figure 256 Vaccinium corym- bosum. West Plain, edge of Cedar Swamp, August 31, 1910. Figure 257 Vaccinium macro- carpon. leathery leaves, 6 millimeters broad. dermal cells are protected by a thin cuticle. are two rows of short palisade cells with large chloro- plasts. The spongy parenchyma is open and of rounded cells. The stomata with their guard cells at the surface are confined to the lower epidermis. A diphotophyll (Fig. 257). Vaccinium pennsylvanicum [Shamong, Upper Plain]. — The early sweet blueberry is a dwarf plant with green, warty, glabrous stems. The leaves are lanceolate, or oblong, distinctly serrulate, bright green, smooth and shiny on both sides, hairy on the midrib above and below. The epidermal cells are large and thin-walled. The stomata are confined to the lower epidermis with their guard cells level with the surface. There are 2 rows of palisade cells and the spongy parenchyma is rather compact in the leaf of the plant from the Upper Plain, more open in Figure 258 Vaccinium pennsyl- vanicum. West Plain, June 28, 1910. 282 VEGETATION OF THE NEW JERSEY PINE-BARRENS Figure 259 Vaccinium vacillans. East Plain, August 31, 1910. the leaf from the pine-barrens at Shamong. A diphotophyll (Fig. 258). Vaccinium vacillans [Shamong and Lower Plain]. — The late low blueberry is a small shrub, 3 to 9 decimeters tall. The leaves are glabrous, obovate or oval, very pale, or dull, glaucous, at least underneath. The upper epidermis of thin-walled cells is covered with a thin cuticle. The lower epidermis of smaller, thin-walled cells is pierced by stomata, the guard cells of which are level with the surface. There are two rows of palisade cells in the leaves from Shamong and the Lower Plain. A diphotophyll (Fig. 259). Pyxidanthera harhulata. — The flowering-moss is a prostrate, creeping plant, with narrowly oblanceo- late, awl-pointed leaves, 3 to 8 millimeters long, and of two colors, brownish and green. The epi- dermal cells of both lower and upper leaf surfaces have a very thick outer wall. The stomata are con- fined to the fiat surface, while the curved surface is without stomata, and here the several rows of palisade cells are located of the stomata are at the surface. A diphotophyll, approaching a spongophyll (Fig. 260). Melampyrum linear e { = M. americanum). — The cow-wheat is an annual plant with opposite, linear- lanceolate leaves. The upper and lower epidermal cells are large, the upper exceptionally so, some of them being modified into short, canine-tooth-like hairs. The stomata have large projecting guard cells. There are two rows of pali- sade cells and the spongy parenchyma cells are large. A diphotophyll (Fig. 261). Chrysopsis falcata [Lakehurst]. — The sickle- leaved, golden aster is a very woolly perennial herb, i to 3 decimeters tall, with linear leaves. The cuticle is developed faintly and the epider- mal cells of the upper and lower sides are large and thin-walled. There are 3 rows of palisade cells and the stomata developed on the lower Figure 260 Pyxidanthera barbu- lata (Shade), March 25, 1910. The guard cells Figure 261 Melampyrum lineare. East Plain, August 31, 1910. Figure 262 Chrysopsis falcata, Lakehurst. VEGETATION OF THE NEW JERSEY PINE-BARRENS 283 side of the leaf have their guard cells level with the general surface. A diphotophyll (Fig. 262). Chrysopsis mariana [Shamong]. — The golden aster is a perennial plant, silky, with long, weak hairs. The oblong leaves have thin-walled epidermal cells and the stomata on the lower and upper surfaces have slightly protruding guard cells. There are two rows of palisade cells, more pronounced in the leaf from the Lower Plain. A diphotophyll (Fig. 263). Eupatoriiim verhenaefolium [Shamong]. — This plant is roughish pubescent. The leaves are ovate-oblong to ovate-lanceolate, slightly three-nerved and coarsely toothed. The epidermal cells are large and thin- walled, bearing tooth-like multicellular, short, rigid hairs on both sides. A diphotophyll. Liatris graminifolia [Shamong]. — This species of blazing-star has linear, one-nerved leaves, borne on a stem 3 to 9 decimeters tall. The upper and lower large epidermal cells are without a cuticle, and both surfaces display stomata with slightly depressed guard cells. There is a single row of palisade cells and the loose paren- chyma cells are rounded. Vesicular hairs are found in slight pits, or depressions on both the upper and lower epidermal surfaces. A dipho- tophyll. Solidago sirida [Shamong]. — The wand-like goldenrod is a smooth perennial with small, entire, appressed, lanceolate-oblong, thickish leaves. The epidermal cells on both sides are large and rounded with stomata on both sur- faces, the guard cells slightly projecting. The palisade parenchyma occurs on both sides in a loose, somewhat irregular arrangement of the cells. The spongy parenchyma occupies a central position. Short canine-tooth-like hairs are found on both surfaces. A dipho- tophyll (Fig. 264). Figure 263 Chrysopsis mariana, East Plain, August 31, 19 10. Synopsis of Leaf Structure An analysis of the leaf structures of Bermuda strand plants, of New Jersey strand plants, of New Jersey salt marsh plants, as well as those of the pine-barren region, brings out some interesting facts about the relative abundance of the different types of leaf structure (Fig. 265). Figure 264 Solidago stricta, Shamong, August 29, 191 1. 284 VEGETATION OF THE NEW JERSEY PINE-BARRENS Such a comparison is important, because it enables us to point in part to this or that structure as pecuHar to plants growing under different en- vironmental conditions. The plant names are omitted, and only a simple enumeration is attempted. ENUMERATION OF LEAF STRUCTURE Special Leaf Structure 1. Cuticle Present 2. Thick Epidermis 3. Hypodermis Present 4. Single Row Palisade 5. Twoor more Palisade Rows 6. Guard Cells Depressed .... 7. Leaves Hairy 8. Leaf Surface Papillate . . . . 9. Leathery Leaf 10. Wiry Leaf 11. Latex Present 12. Crystals Present 13. Idioblasts 14. Diphotophyll 15. Diplophyll 16. Staurophyll 17. Spongophyll 18. Succulent Leaf Ber- muda Dune Plants 10 5 New- Jersey Dune Plants 8 7 I 5 2 I 4 I 10 3 2 2 3 II 55 Plants of Pine-Barren New Jersey Salt- Marsh Plants Pine Forest 18 3 24 10 '7 5 >3 5 I 9 29 3 4 I White- Cedar Swamp Plains It will be noted in the accompanying table that the plants examined from the six different localities are diphotophylls, and this is true especially of the plants that grow in the pine forest. The diplophyll structure, which is occasioned probably by illumination on both leaf surfaces, is a prominent feature of the dune plants of Bermuda and New Jersey, as is also the staurophyll. The spongophyll is found in dune plants and salt-marsh plants, while it is absent almost entirely (i case) in the plants of the pine-barrens. Hairy leaves are frequent in pine- barren plants, as also those with a cuticle. Two or more rows of pali- sade cells are more common in pine-barren plants than a single row, while in the dune plants a single row of cells is more frequent than a double row, or over. More dune plants have a thick epidermis than do pine-barren plants. The papillate leaf surface is a feature of the pine- barren plants and leaf succulency is a dune and salt-marsh characteristic. Enough has been said to show that the table is instructive and enables VEGETATION OF THE NEW JERSEY PINE-BARRENS 285 US to contrast the influence of the different ecologic factors upon the structure of the plants concerned (Fig. 265). Figure 265 Comparative frequency of various leaf structures of plants of (A) Bermuda dunes; (B) New Jersey dunes; (C) New Jersey salt marsh; (D) New Jersey pines; (E) New Jersey cedar swamp; (F) New Jersey plains; XX indicates absence of structure. CHAPTER XX CONE AND SEED PRODUCTION OF THE PITCH PINE August Renvall* has presented, in a monograph pubHshed at Helsingfors in 1912, the periodic phenomena of the reproduction of the Scotch-pine, Pinus silvestris, at the polar Hmit of trees in Finland. He considers, after a valuable historic introduction, the formation of staminate and ovulate flowers, working out statistically the frequency data, the steril- ity of the trees, and other important details by biometry. The years of cone and seed production also are discussed by Renvall, who uses the statistic method. Coastal. Dune Complex Inland. 3 I Figure 266 Arrangement of pitch-pine cones to show the years of cone production and by the open cones the years of seed discharge. Note that the coastal pine cones are larger than the inland. Following out the suggestions of this important paper I undertook, during the latter part of the summer of 191 2, to apply somewhat similar methods to the study of the pitch pine, but I have omitted the purely mathematic method and I have emphasized that phase of the study which appeals directly to the forester, who is concerned with the repro- duction of the species. Several trees were taken in each locality and a single branch or two on each tree were chosen upon which to make the estimation of cone production (Fig. 266). The localities are given in each enumeration. * Renvall, August: Die periodischer Erscheinungen der Reproduktion der Kiefei anderpolaren Waldgrenze, Fennia 29, No. 4, 1912, pages i-xii, and 154, with colored map. 286 VEGETATION OF THE NEW JERSEY PINE-BARRENS 287 Belmar Park, N. J. — The first tree studied had a circumference measurement of 76 decimeters, or an approximate diameter of 28 deci- meters. A lower, south-bending limb was taken. The numbers after the year indicate the number of cones found on the portion of the branch corresponding to that date. First Limb. — 1912=1; 1911=4; 1910 = 3; 1909=1; 1908 = 2; 1907 = 3; 1906 = 4; 1905 = 1 broken off; 1904 = 2 one broken off; 1903 = 2 broken off ; 1902=1; 1901 = 1; 1900=1; 1899 = 0; 1898 = 0; 1897 = 0; 1896=1 detached and hanging by the bark. Second Limb. — 1912 = 0; 1911=2; 1910 = 4; 1909 = 0; 1908 = 2; 1907 = 2 one broken off; 1906 = 0; 1905=4; 1904 = 0; 1903=1 broken off; 1902 = 2; 1901=2; 1900 = 0; 1899 = 0; 1898= I broken off; 1897 = o; 1896 = 0; 1895=2. The second tree, which was from twenty-five to thirty years old, was chosen in the same locality. 1912=1; 1911=4; 1910 = 2; 1901 = 1 dried up; 1908 = 0; 1907 = 0; 1906 = 2; 1905=2; 1904 = 2 one broken off; 1903 = 0. The third tree was twenty-five to thirty years old and grew in the same locality. 1912 = 2; 1911=6 two in one cluster, four in another cluster separated from each other; 1910 = 2; 1909 = 0; 1908=1; 1907 = i; 1906 = 2; 1905=0. The fourth tree grew in the same locality, but in a place more exposed to the wind and sun. 1912 = 2; 1911=0; 1910 = 3; 1909 = 0; 1908 = 0; 1907 = 2; 1906 = 3; 1905=0. The fifth tree grew in the same locality, but was exposed to the north- east winds from the nearby ocean. 1912 = 2; 1911=0; 1910 = 2; 1909 = 0; 1908=1; 1907=1; 1906=1; 1905 = 1; 1904 = 0; 1903=0. It will be seen from the above enumeration that from 1905 to 1912 practically every year was marked by the production of cones on one of the five trees. The year 1905 was perhaps the worst year for cone production, as only one tree out of five, as evidenced by a single branch, produced cones in that year. Each year from 1900 to 1905 on the first tree was marked by the production of cones. Then the years 1897, 1898, 1899 were bad cone years and 1896 a good cone year. Practically all of the cones on these five trees were open, showing that the winged seeds had been discharged during the late fall of the year of their ma- turity. To determine whether the liberated seeds germinated, or were in a viable condition, which is an important consideration to the forester, the expedient was adopted of cutting down young trees with a saw. The annual rings were counted and the diameter of the tree measured. The years in which viable seeds were produced was ascertained by sub- 288 VEGETATION OF THE NEW JERSEY PINE-BARRENS trading the number of annual rings from the year of Study, 19 12. Many factors are influential in the production of seeds and cones and these are spread over two years, hence the conditions of the year preceding the actual formation of cones and seeds may have been influential. With this explanation the statement given above as to the year in which the ripe cone and ripe seeds were formed is not misleading. Ten trees were thus measured at Belmar Park with the following results: Tree I 2 3 4 5 6 7 8 9 10 Number of Rings 14 14 14 10 10 Diameter 9.0 cm. 8.0 5-5 5-5 3-4 2-5 2.2 2.6 2.0 1.8 Year of Germination i»94 1898 1898 1898 1902 1902 1904 1904 1905 1908 It will be seen, as far as the estimation of the 10 young trees indicates, that there were viable pine seeds produced, as evidenced by those which germinated in the years 1894, 1898, 1902, 1904, 1905, and i 908, although, as previously shown, the pitch-pine trees produced seed every year (Fig. 268), but the conditions suitable for germination may have been absent or the seed may not have been fertilized, and other suitable factors may have been absent during the years unaccounted for in the above table The second locality was chosen in the dune complex area at Spring Lake, where the bases of the trees were surrounded with the marram- grass, Ammophila arenaria, and clumps of the wax-myrtle, Myrica carolinensis. The first tree was about 2 meters (6 feet) tall, of a spreading, wind- swept form, with 18 rings, and grew near the inner edge of the dune complex. The diameter of the tree was 70 centimeters. The count of the cones was made on a branch of the lee side. 1912=1; 1911=2; 1910 = 2 tightly closed; 1909 = 0; 1908=1 tightly closed; 1907 = 0; 1906=1; 1905=0. The second tree was a low-spreading specimen near the outer edge of the dune complex at Spring Lake. The first branch showed the fol- lowing: 1912=1; 1911=2; 1910 = 2; 1909=1. The lower, cone-bear- ing branch of this tree, well protected by the dense crown of the tree. VEGETATION OF THE NEW JERSEY PINE-BARRENS 289 showed: 1912=0; 1911=0; 1910 = 0; 1909 = 0; 1908 = 0; 1907 = 0; 1906 = 2 tightly closed. The third tree grew at the extreme outer edge of the dune complex. It was a tree i meter tall with yellowish-brown leaves. The whole tree had only 2 cones both unopened, and by a count of annual rings, it was 9 years old. The whole appearance of the tree betokened stress of growth conditions. First Limb. — 1912 = 0; 1911=0; 1910 = 0; 1909=1 tightly closed. Second Limb. — 1912=0; 1911=0; 1910= i tightly closed. The fourth tree, over 3 meters tall, grew in the thicket back of the dune complex. 1912=1; 1911=0; 1910 = 0; 1909 = 2 green; 1908 = 0; 1907=1 tightly closed. The fifth tree in this locality grew at the outer edge of the thicket and was 10 feet tall. A branch on the protected side was studied. 1912 = 5 in one whorl, 3 in another whorl lower down; 1911=3; 1910 = 0; 1909 = 2 tightly closed; 1908= i tightly closed. Two other trees were cut down in this thicket, one showed 9 annual rings and, therefore, began its growth in 1903, reaching a stem diameter of 7 centimeters; the other showed 13 rings and began its growth in 1899 (Fig. 266). A study of these dune trees show that the cones were not produced every year, and even in those years when cones were developed their production amounted to nothing in the regeneration of the forest, as long as they remained tightly closed. It shows that the wind-swept trees at the outer part of the dune complex are at the edge of their natural range as long as the present conditions control, because these trees rarely produce cones, and those that are formed, never open to let their seeds fall out. The advance of the forest into the dune complex as the pioneer trees prepare the way for those that follow, therefore, depends on the seeds which are produced on trees that grow some dis- tance back from the outer edge of the tree line of the dune complex.* These winged seeds are carried to the tension line between the dunes and the forest by the currents of wind that blow. Not every year wit- nesses the start of a new tree. In the area studied new trees began their growth in the years 1894, 1899, and 1903. To determine the cone production under normal conditions of forest growth a locality was chosen at Spring Lake, about i kilometer (K mile) inland. The pine wood opposite to the "Plantation" in Spring Lake was chosen, where there was a uniform stand of tall pine trees with * Cf. observations by Robert F. Griggs on the encroachment of the Sitka spruce on the tundra at the northern limit of forest on Kodiak island, Alaska, Bull. Torr. Bot. Club, July, 1914. •9 290 VEGETATION OF THE NEW JERSEY PINE-BARRENS at least forty years of growth. The heart-wood covered 6 annual rings. A number of young trees were cut down at the southern edge of the woods. The following data were obtained: Tree Number of Rings Diameter in Centi- meters Ye.\r of Germination I 14 12 12 10 7 5 6.0 . 2.5 Suppressed. 3-5 4.8 1.8 1-3 1898 2 1900 a 1900 A 1902 e; 1905 6 1907 The largest young trees, averaging from fifteen to twenty years, grew on the west, or lee side, of the taller trees. The younger trees form a fringe to the woods, because they start best in well-lighted situations, as the pitch pine is intolerant of shade. Tree No. 2 shows this in the small diameter, as compared with a tree fourteen years old growing in a sunny position. Nearly all of the young trees beneath the larger ones were similarly suppressed. There is very little typic pine-barren under- growth in this particular grove, because the taller pine trees probably grew up in an open field forty to fifty years ago. A study was made of a tall pitch-pine tree some distance west of the "Plantation." The circumference of this tree was 9.7 decimeters, which would make its diameter about 3.2 decimeters. 1912 = 2; 1911=4 green unopened; 1910 = 2; 1909 = 3; 1908 = 0; 1907 = 2; 1906 = 3; 1905=2; 1904 = 2; 1903=0; 1902 = 0; 1901 = 1. Another locality was chosen 6 kilometers (4 miles) inland, i. c, 3 kilometers (2 miles) west of Point Pleasant. It was found that there was no uniformity in the production of cones on different branches. A tree twenty-six years old was found to be 7.6 meters tall. The diam- eter, 6 decimeters above the ground, was found to be 1.2 decimeters. Rapid growth was made in the first fifteen years, and in the later eleven years of growth the rings were small and difficult to see. The year 1911 was an exceptionally good year in this locality, as indicated by green cones on nearly all of the pine trees for the growth of that year, but, as emphasized previously, we must presume that pollination was successful the year before. The tree studied was cut down in order to get at the cones (Fig. 266). First Limb (topmost).— 1912 = 2; 1911 = 1; 1910 = 2; 1909 = 0; 1908 = 0; 1907=1; 1906=1; 1905=0. VEGETATION OF THE NEW JERSEY PINE-BARRENS 291 Second Limb. — 1912=1; 1911=2; 1910=1; 1909 = 0; 1908 = 3; 1907=1; 1906 = 2; 1905 = 1 with the cone partially imbedded in the bark. A number (3) of small trees were cut down. Number of Tree Number of Rings Height in Meters Diameter in Centimeters Year of Germina- tion I . . . 18 •5 10 1.82 1.52 1. 21 • 3-3 2-5 2.8 1894 1897 1902 2 3 A study of a tree in the heart of the pine-barrens on the Speedwell Road outside of Shamong (Chatsworth), N. J., on November 13, 1912, resulted in the following: 1912 = 0; 1911=3; 1910 = 0; 1909=1; 1 908 = I ; 1 907 = 2 ; 1 906 = 3 ; 1 905 = 2 ; 1 904 = 3 ; 1 903 = 2 ; 1 902 = 3 . On another smaller tree cones were found for the year 1912. It will be noted that in the heart of the pine-barrens the production of cones that open finally to discharge their seeds is the normal course of events. If we take the photograph of the museum preparation illustrating the pro- duction of pine cones under various environmental conditions, we find that the pine cones of the coastal trees are larger than those produced in the interior, as far as the observations of the writer go. I would account tentatively for this difference in size as due to a larger amount of mois- ture in the air of the coastal pine-barrens, as contrasted with the drier air inland. The conditions of the interior, however, are such as to produce smaller cones (Fig. 266). The following conclusions have been reached from such a statistic study of the seed and cone production of the pitch-pine tree in the New Jersey pine-barrens. The regeneration of the pitch pine, Pinus rigida, depends on the following factors which are operative over two years: First: The formation of viable seeds. This is conditioned on: {a) Proper pollination. {h) Certain fertilization, (c) Proper development of the embryo. {d) Absence of seed-eating insects and rodents. Second: On the opening of the pine cones at the end of the second year, when they reach maturity. This normally takes place in the pine- barrens of New Jersey in the Indian summer days of November. Third: On a suitable seed-bed. Fourth: On climatic conditions suitable for germination. Fifth: On plenty of light. 292 VEGETATION OF THE NEW JERSEY PINE-BARRENS Sixth: On freedom from too much competition. Seventh: Absence of fire for a few years after germination. The absolute success of the estabhshment of a pitch-pine tree is de- termined best by an actual study of young trees in all stages of growth toward maturity. Law of Cone and Seed Production in the Pitch Pine in the Pine- Barren Region of New 'Jersey Cone production in the pine-barren region of New Jersey (coastal and inland) is an annual event, followed by the opening of the cones in the late fall of the year after their inception, or later, and the discharge of viable, winged seeds. The omission of this event is a local phenomenon and not of general occurrence in any one year. Natural annual regenera- tion is assured if the factors described above are favorable. Vivipary in the Acorns of Quercus Marylandica On September 5, 191 2, at Spring Lake, New Jersey, an oak tree of this species was found with a viviparous acorn. This condition was induced probably by the heavy rains, followed by mists and fogs, that had pre- vailed for a few days previously. An examination of this acorn showed that the embryo had swollen sufficiently to crack open the acorn cover- ing, which had started to glaze. This splitting of the acorn shell revealed the presence of a small lenticular acorn at the base and inside the margin of the cupule. This vivipary of the acorns has not been studied exhaustively in America, but in Europe it has been studied in Quercus robur. Guppy* states that this is exhibited, not only in the occasional germination of the fruits on the tree, but in the actual stages of growth of the seed within its shell before maturity is reached. The steady growth of the seed on the tree long after the pericarp, or shell, has begun to dry is noteworthy. The vital connection with the parent plant is maintained by the attachment of the base of the fruit to its cupule. When the acorn begins to brown this attachment to the cupule begins to loosen, the result evidently of the drying of the pericarp, or shell. In the case of the viviparous acorn, it falls to the ground and usually dies, but it must happen frequently in moist mild weather that it continues the growth commenced on the tree, and if covered by pro- tecting leaves, it may survive to the next spring and grow into an oak tree. Guppy suggests in connection with this viviparous habit, that if acorns are taken before they enter the rest period, that is while the peri- * Guppy, H. B.: Studies in Seeds and Fruits, 310 and 432. VEGETATION OF THE NEW JERSEY PINE-BARRENS 293 carp is still green but the embryo is mature, they can be induced to keep up an uninterrupted growth without entering the rest period. This suggestion was tested with a number of acorns of species of oak, viz., Quercus alba, Q. marylandica, Q. prinus. Acorns of these three species were planted in a box filled with sphagnum. The planting was done on September 12, 191 2, and the results recorded on October. 22 1 91 2. Two series of acorns were sown. One set had a portion of the shell removed, exposing the embryo. The other set was planted with an uninjured shell covering. There was a slight advantage in the rate of germination of the cut acorns, as contrasted with the uncut. Practically all of the green acorns of the chestnut oak, Quercus prinus, the white oak, Q. alba, and the black-jack oak, Q. mary- landica, germinated (Fig. 267). Sachs* and other botanists have maintained that, even under the most favorable conditions of vegetation, dormant periods occur in the course of the life of the plant. Under circumstances when the plant would be in a condition to grow most vigor- ously, because it is provided with reserve materials, water and oxy- gen are at its disposal and a suf- ficiently high temperature might be expected to call forth the in- ternal activities, yet every ex- ternally perceptible vital motion nevertheless ceases, and it is only after some months of rest that the growth commences anew, and this frequently under circumstances which appear far less favorable— especially at a conspicuously lower temperature. The experiments with acorns detailed above and experiments performed by Guppyt with the seeds of Iris pseudacorus, Vicia sepium, Arenaria peploides and Quercus robur show that a rest period is not essential for the germina- tion of acorns and other seeds, but that by taking immature acorns^ whose embryo has not ceased to grow, and planting them the period of * Sachs, Julius: The Physiology of Plants: 350. t GuppY, H. B.: Studies in Seeds and Fruits: 421. Figure 267 Stages in vivipary of Quercus marylandica A, Acorn of Quercus marylandica. Acorn of Quercus marylandica dis- sected. Embryo of Quercus marylandica. First stage of vivi- pary in acorn of Quercus mary- landica. E, F, G, Later stages of vivipary. H, I, Stages in germi- nation of green acorns of Quercus marylandica. J , Stage in germination of green acorn of Quercus alba. K, Details of such germination. 294 VEGETATION OF THE NEW JERSEY PINE-BARRENS growth is maintained without cessation, or a rest period, and the result is the elongation and growth of the embryo into a young seedling plant, as fully demonstrated in the figures. The germinative capacity of so- called unripe seeds does not seem to have been appreciated by foresters and gardeners, who layer their tree seeds in boxes of sand kept slightly moist and stored in a cool place over the winter. The acorns can be planted while green and carried over the winter in the greenhouse in the actively growing condition and in the spring, they can be planted in the open (Fig. 267). CHAPTER XXI NOTES ON A FEW INSECT GALLS OF THE PINE- BARRENS Modern ecology has concerned itself with the habitat relationships of plants, and no more interesting field presents itself than the association of insects and plants of such a character that a gall, or cecidium, is the product.* The cecidologist is an ecologist, or biologist, who has chosen a narrow field of study, that of the galls. There are, he tells us, various kinds of cecidia caused by specific organisms, such as insects, nematodes, fungi and bacteria. We are concerned in this monograph only with insect galls. An insect gall is an abnormal growth of plant tissue produced by an insect which inserts an egg into the plant tissue from which hatches out a larva which is the exciting cause of the gall formation. Adler has shown that there was no foundation for supposing that the gall mother injected any irritating secretion whatever, and Beyerinck proved that the fluid ejected by the gall-fly is without taste or smell, and abso- lutely unirritating if injected under the skin. Both of these authors show that it is not in the gall-mother, but in the larva, that we must seek for the cause of the gall growth; and that it is the nature of the salivary secretion, and the manner of feeding of the larva, peculiarities inherited by each species which give the characteristic structure to each specific kind of gall. Mel. T. Cook f has shown that "the morphologic char- acter of the gall depends upon the genus of the insect producing it, rather than upon the plant on which it is produced." The necessity for the continuance of the excitation during the whole period of gall growth is shown by its cessation when the larva has been destroyed by parasites. * Consult Beutenmuller, William: Catalogue of Gall Producing Insects found within Fifty Miles of New York City. Bull. Amer. Mus. Nat. Hist., iv: 245-278; Adler, H.: Alternating Generations: A Biological Study of Oak Galls and Gall Flies, 1894; Connold, Edward T.: British Vegetable Galls, 1902; Beutenmuller, William: insect Galls of the Vicinity of New York. Guide Leaflet, No. 16, Amer. Mus. Journ., iv, No. 4, October, 1904. fCooK, Mel. T.: Galls and Insects Producing Them. Ohio Naturalist, 11: 263-278, 1901; Cecidology in America. Bot.Gaz.,49: 219-222, March, 19 10; Mayr,Gustav L.: Mittel-Europaischer Eichen-Gallen in Wort und Bild, 1907; Kuster, Dr. Ernst: Die Gallen der Pflanzen, 191 1. 295 296 VEGETATION OF THE NEW JERSEY PINE-BARRENS The association of the insect and host plant is of ecologic interest. The fact that a specific larva can produce a specific gall of a definite shape, size, color, and longevity is one of the most remarkable in the whole range of biology. That out of elements so diverse in nature as the living cells of the host plants and the larval body of an insect, can result a gall of great beauty at times and always the same in morphologic structure, if produced on a given plant by a given insect, is remarkable. A few insect galls were collected about 3 kilometers (2 miles) west of Point Pleasant in September, 1 91 2. These galls with one excep- tion were identified by Professor P. P. Calvert, of the University of Pennsylvania, and one of them by Dr. William Beutenmiiller, of the American Museum of Natural History (Fig. 268). On the white oak, Quercus alba, was found a large gall, the oak- seed gall, due to Andricus semi- nator Harris. This gall, which is found on the twigs of the white oak in June, is composed of a woolly substance, and is irregu- larly rounded, about 4 centi- meters in diameter. Inside are numerous seed-like bodies adher- ing round the twig and very much , resembling seeds, hence the com- mon name. The gall is pure white, or white tinged with red, but by September it assumes a rusty-brown shade. Later it breaks to pieces and drops off the twig. The insect is hymenopterous, belonging to the Cynipidae (Fig. 268A). The scarlet oak, Quercus coccinea, had a number of different galls upon it. The oak, or May-apple, the largest of these galls, is due to an hymenopterous insect, Amphibolips confluentus Harris, which attacks the oak leaves. The gall is large, globular and smooth outside. The interior is filled with a spongy substance surrounding a central, hard, woody kernel containing the larval cell. The gall is about 2.5 centi- meters in diameter and when fresh is pale-green, soft and succulent with Figure 268 Insect galls on pine-barren trees. A, Oak seed gall, An- D, Oak Apple, Amphi- dricus seminator bolips confluentus Harr. Harr. B, Amphibolips coelebs E, Cecidomyia pisum, O. S. Fitch. C, Holcaspis centricola F, Coccid. O. S. VEGETATION OF THE NEW JERSEY PINE-BARRENS 297 whitish contents. The shell becomes light-brown in September and brittle. Another uncommon gall, which lives attached to a vein on the underside of the scarlet oak leaf, is the oak spindle gall, Amphibolips coelebs Osten Sacken (Fig. 268, B). This gall is elongated, spindle- shaped, at first soft and green, later becoming umber-brown and brittle. The central kernel is held in place by radiating fibers. The scarlet oak specimen yielded a small, spheric gall, determined by Dr. William Beutenmuller to be caused by Cecidomyia pisum Fitch, a dipterous insect. This little gall distributed plentifully over the leaf surface which is roughened by it, is deep reddish-brown in color, and woody. it is about the size of a morning-glory seed. A coccid was found also attached to the twigs in the axils of the leaves. On the leaves of the post oak, Quercus stellata, was found a large, spheric gall due to Holcaspis centricola Osten Sacken, a hymenopterous insect. This gall is about 2 centimeters in diameter, and in September is of a light yellow-brown color. The shell is smooth and brittle. In- ternally there are radiating fibers attaching a central body to the inner wall of the gall (Fig. 268, C). Four of the above gall insects, viz., Andricus seminator, Amphibolips coelebs, A. confluentus, Holcaspis centricola, are hymenopterous, be- longing to the family Cynipidae, the " gall-wasps," or "gall-flies." The ovipositor of the adult insect is coiled partly within the abdomen, which is dilated and enlarged posteriorly, closely joined to the thorax, but not sessile. The life-cycle is often complicated, and parthenogenesis is of frequent occurrence. In some species the walls, perhaps, have been eliminated, while in others there is an alternation of generations, one having both sexes normally present, while in the other the females only occur. Cecidomyia pisum is a dipterous insect, included in the family Cecidomyiidae, the "gall-gnats," or "gall-midges," which are fragile in appearance and slow in flight. The larvae are small, legless grubs, bluntly pointed at both ends, often with a chitinous process, known as a breast-bone, on the under side, near the anterior end.* A complete list of the gall insects of New Jersey is given in the volume on insects, Annual Report New Jersey State Museum, 1909. The interested bot- anist is referred to that publication for additional details about the pine- barren gall insects. * Consult Report of the New Jersey State Museum. The Insects of New Jersey, 1909: 595-604; 725-734. CHAPTER XXII PINE-BARREN PLANTS FROM AN EVOLUTIONARY VIEWPOINT Our study of the vegetation of the pine-barrens of New Jersey would not be complete without reference to the broader principles which under- lie all such ecologic studies, namely, questions of evolution. Many of the evolutionary problems in Europe and America have been approached by a study of cultivated plants. While cultivated plants are suitable for the investigation of many phases of evolution, yet they are not suited so perfectly for this purpose as the wild plants which exist in virgin plant formations. For this reason the pine-barren vegetation of New Jersey, which exists practically in undisturbed condition in its original regional surroundings, affords unusual opportunities for research upon plants in a state of nature. De Vries has emphasized the importance of the experimental treatment of the species question, but in the im- possibility of studying all species experimentally the ecologist and taxonomist, with different ends in view, can contribute much material which will be of lasting value. The taxonomist is prone to make new species, because his object is to describe a plant so that it may be identi- fied readily, and this, from the very nature of the case, demands a more or less artificial classification. The species of the taxonomist, as L. Cockayne* has proved abundantly, are frequently ideas merely and not living entities. The ecologist, on the other hand, emphasizes the in- fluence of the environment in modifying the form of the species, and he finds that the descriptions of the taxonomist do not always fit the plant, even with an elastic interpretation of the specific characteristics of the plant. With him a specific name is a means, rather than an end, be- cause in his description of vegetation the scientific names are useful as a means of describing intelligibly the associated plants of a particular plant formation and the epharmonic structures of these plants. He recognizes the necessity of the substitution of elementary species, as the raw material for the evolutionary process, rather than the Linnaean historic or taxonomic species. * Cockayne, L.: Observations concerning Evolution, derived from Ecological Studies in New Zealand. Trans. New Zealand Instit., xliv: i — 50, 1912. 298 VEGETATION OF THE NEW JERSEY PINE-BARRENS 299 Figure 269 The spurge, Euphorbia ipecacuanhae, illus- trating in its various forms the origin of elementary species. Elementary Species. — Elementary species are physiologic concep- tions. They are forms whose characters, whether large or small, are heritable, and thus they become of phylogenetic importance. Among the pine-barren plants no one plant possesses more interest from the standpoint of elemen- tary species, than Euphorbia ipecacuanhae,* in which Pro- fessor de Vries took the greatest interest, when the writer showed him the plant some years ago in its natural habitat in New Jersey. This taxonomic species consists of distinct forms that merit specific rank as elementary species. The leaves in the different types range in shape from linear-lanceolate to broadly elliptic, and in color from green to deep reddish-brown, and these leaf and color characters are heritable (Fig. 269). Thecowbane, Oxypolis rigidior, and its variety longifolia, prob- ably represent two elementary species. The variety has narrow leaf segments, rarely over 4 to 5 mm. wide. The margin of the segments are generally entire, but the narrowest are sometimes lobed (Fig. 270). The writer has been interested for some time in Vaccinium co- rymbosum and V. atrococcum. Both are tall blueberries growing in wet thickets. The first shrub flowers in May, when the leaves are partly expanded and the fruit is black with a bloom. Vacci- nium atrococcum flowers earlier Figure 270 The cowbane, Oxypolis rigidior, in its sev- eral varieties which are probably elemen- tary species. The lower figure represents the variety longifolia. than the preceding, and its leaf development follows usually the open- ing of its flowers. The fruit is black without bloom. Two other * DE Vries, Hugo: The Mutation Theory [Engl, transl.], 11: 605, 1910. 300 VEGETATION OF THE NEW JERSEY PINE-BARRENS taxonomic species are distinguished, viz., Vaccinium virgatum Ait. and V. caesariense Mackenzie. According to Mackenzie, there are apparently three forms of tall blueberry native to the coastal plain of New Jersey, viz.: (i) a form with finely serrate leaves (virgatum), somewhat pubescent below, and restricted to the pine-barrens; (2) a form with entire leaves, partially pubescent below, particularly the veins (corymbosum) ; and (3) an entire-leaved, absolutely glabrous form (caesariense). Perhaps we should think of all these four forms as ele- mentary species, viz., Vaccinium atrococcum, caesariense, corymbosum, virgatum. The small-flowered dogbane, Apocynum medium Greene, as a taxo- nomic species, includes probably two growth forms which have been segregated from it, namely, A. Milleri Britton and A. urceolifer G. S. Miller. These are prob- ably all elementary species. Variation. — Darwin empha- sized the study of individual differences of organisms in his discussion of variation. He con- sidered that they formed the material upon which natural selection could act, and hence they accumulate. De Vries and others deny that these fluctuat- ing characters are accumulated indefinitely, and as far as the writer's study of plants enables him to form an opinion, he agrees with the position of de Vries. We can, by statistic methods, fix the limits of such fluctuations, as they oscillate around an average. A number of interesting cases of fluctuating variability occur among the pine-barren plants of New Jersey. The pitch pine, Pinus rigida, shows a great diversity of tree forms, which have been described in a former section. It is difficult to determine which is the normal form of the tree, although in fact no one has any difficulty in pointing out a pitch-pine tree when he sees one. Such fluctuations show how difficult it is to give a description of a tree which will be comprehensive enough to include all of the different growth forms (Fig. 271). Figure 271 Different forms of branches from pine trees growing in different localities. Left, pine forest at Belmar; Second, branch from a protected tree, sea dune thicket; Third, branch from a small tree in dune complex; Right, branch from exposed side of the same tree from which No. 3 branch came. VEGETATION OF THE NEW JERSEY PINE-BARRENS 301 The common arrowhead, Sagittaria longirostra, of the pine-barren region, shows leaves that vary from narrowly linear to broadly hastate, though the narrow type of leaf is more frequent (Fig. 272) . Panicum Commonsonianum is a plentiful grass throughout the pine-barrens, first described by Ashe. Related to it is another slightly different grass, distinguished by Nash as the va- riety Addisonii. Between these two Figure 272 Leaf variation of Sa A-B, Leaves from a single plant col- lected by S. S. Van Pelt at Tom's River, July ig, 1906. Narrow leaf an early juvenile form at the out- side of the rosette. C, Leaf of plant col- lected by Bayard Long at Lake- hurst, August 8, 1908. D, Leaf of plant col- lected by J. H. Grove at New Egypt, August 14, 1907. E-F, Leaves of a single plant collected by Bayard Long at Dover Forge, Sep- tember 9, 1907. Narrow leaf from outer part of ro- sette an early ju- venile form. gittana longirostra. G-H, Leaves of a single plant collected by S. S. Van Pelt, near Job's Bridge , between Ham- monton and Bat- sto, August 6,1905. L Leaf of a plant from Parkdale collected by S. S. Van Pelt , August 17, 1905. J, Leaf from Lakehurst plant collected August 8, 1908, by Bayard Long. K, Leaf of a Pasadena plant collected by Bayard Long on July 24, 1908. L, Leaf of a plant col- lected by Bayard Long at Lake- hurst, August 8, 1908. Figure 273 Leaf variation of Peitandra virginica. A-B, Leaves of a single D, Leaf of a plant col- plant as denoted lected by Alexan- by connecting der Mac Elwee dotted line, col- at Forked River, lected by Bayard June 22, 1895. Long at Mana- E, Leaf of a plant col- hawkin on Sep- lected by Bayard tember 16, 1910. Long along Hospi- C, Leaf of a plant col- tality Branch at lected by J. H. Folsom on June Grove at New 10, igir. Egypt, June 22, F, Spathe of the plant . 1906. forms are a great number of intergradations. The green arrow arum, Peitandra virginica (Fig. 273), shows also a great variety of leaf forms, 302 VEGETATION OF THE NEW JERSEY PINE-BARRENS Figure 274 Leaf variation of pickerel weed, Pontederia cordata. A-B, Leaves from a plant collected between D, A whole plant collected by A. N. Leeds at Tuckerton and Atsion by C. F, Saunders Mays Landing, August 20, 1893. and W. N. Clute, July 3 to 6, 1899. E, Leaf from a plant collected at the mouth of C, Leaf from pickerel weed found at Toms River, Hamilton Creek, August 21, 19 10, by Bay- June 2, 1895, by Alexander Mac Elwee. ard Long. VEGETATION OF THE NEW JERSEY PINE-BARRENS 303 Figure 275 Variation in the leaves of Smiiax tamnifolia. as also the pickerel weed, Pontederia cordata, which has a series of leaf forms from broad to narrow, the narrow forms being distinguished as the variety angustifolia by Pursh (Fig. 274). The number of flowers varies considerably in the turk's-cap lily, Lilium superbum, according to locality. Usually only a single flower is found on plants growing in the pine-barren swamps, while in richer swamps of West Jersey it bears a large cluster of flowers, as many as fifteen to twenty. The colic-root, Aletris farinosa, exists in a form with short leaves and more nearly spheric flowers found near the plains. Smiiax tamnifolia (Fig. 275), Quercus falcata (Fig. 276), show the greatest diversity of leaf forms, which are cases of fluctuating variability, and in a less pro- nounced way we note the same in the leaves of Quercus alba, Q. ilicifolia, Q. marylandica, Q. stellata, and Q. velutina. (See Figs. 277 and 278.) The forms of the spatterdock (Nymphaea) in the pine region of New Jersey vary considerably. Stone believes that Nymphaea rubrodisca is referable to N. variegata; as also those referred to N. micro- phylla, which is connected with N. variegata by a full series of intermediates. The flower color of Lupinus perennis shows great fluctuations from deep-purple to white. As intermediate colors exist between these two colors, we must classify these color variations as of the fluctuating sort. The meadow-beauty, Rhexia virginica, shows varia- tions in flower color to deep magenta, but never the pale pink of R. mariana. Mutations. — The origin of new species in the sense of de Vries by mutation can be proved only by experiment under which the heredity of the new character may be proved. In a state of nature, therefore, Figure 276 Variation in the leaves of Spanish oak Quercus falcata (= Q. triloba). 304 VEGETATION OF THE NEW JERSEY PINE-BARRENS we can infer only that such sports or mutants have arisen. The new forms are distinct from each other and do not show intermediate types, such as we find in variations of the fluctuating type. In surveying the plants of the pine-barrens it seems that the following cases are muta- tions. Pogonia ophioglossoides is a bog orchid, usually with a single, rose- colored flower. Occasionally mutants are found with white flowers. The water lily, Castalia odorata, has beautiful, creamy white flowers. Sometimes a pink color suffuses the whole flower. Some authors call Figure 277 Leaf variations of oak, Quercus marylandica. Figure 278 Leaf variations of Quercus stellata and Q. iiicifoiia. this the variety rosea. Polygala cruciata has flowers that are normally the color of red clover, that is, rose-colored, but sometimes as a mutation we fmd plants with greenish-purple flowers, and this statement is true for Polygala viridescens, with rose-colored flowers. Occasional forms have green flowers. These two color forms were described originally as different species. The calico-bush, Kalmia latifolia, has masses of flowers of deep pink, but white-flowered bushes are not infrequent. The bird-foot violet, Viola lineariloba, has large, lilac-purple flowers. As a mutant of this species, the true Viola pedata, we have flowers in which the two posterior lateral petals are a dark velvety purple. The broom crowberry of the plains, Corema Conradii, exists in two color VEGETATION OF THE NEW JERSEY PINE-BARRENS 3O5 forms, deep green and reddish-brown. These forms are both found in adjacent patches in the bright sunHght and are probably mutants. As a mutant of the widely distributed red-maple, Acer rubrum, we have the variety tridens of Wood, or Acer carolinianum of Britton. The leaves of the mutants are small, with three short lobes, the middle lobe being triangular. Epharmony. — Epharmony is the harmony between the structure of a plant and the external factors. Vesque* says: "Tous les organes de la plante peuvent s'adopter au mileu incerte ou anime qui les entoure, mais a des degres divers, et c'est precisement sur cette inegalite que re- pose la subordination des caracteres; mais au mileu de tous ces organes il y en a dont la nature depend 'uniquement de I'adaptation,' savoir la structure anatomique des organes vegetatifs en tant qu'elle est en relation directe avec I'air, le sol et I'eau, c'est ce qui je propose d'appeler I'epharmonie." Plasticity of Species. — Nothing has been brought out more clearly by ecologic studies in the pine-barrens of New Jersey than the extreme plasticity of many species and structures, and their response to a change of environment. This fact is so evident that the idea of normal loses its value. For example, the black-jack oak, Quercus marylandica, may exist in the pine-barrens as a tree and as a shrub. The pitch pine, Pinus rigida, we have seen has a great variety of growth forms and its plasticity is manifested also in the size of its cones. Response to Ecologic Factors. — Warming, in his "Oecology of Plants," 1909: I and 8i, has summed up our knowledge about these matters, so that only a few examples are concerned. (a) Soil Pyxidanthera barbulata, as we have seen, exists in three forms which have been conditioned on the character of the upper layers of the forest soil. The wandering stems of one form of this plant are due to soil condi- tions. Two other forms of this plant occur in the pine-barren region of New Jersey, as forms due to the influence of the substratum as previously noted. (h) Light Quite a number of plants show two distinct forms which grow in the shade and in the light. The pitcher-plant, Sarracenia purpurea, has green ascidia when growing in deep, shady white-cedar swamps. In open sunny places it is generally a deep shade of reddish-brown. The * Ann. Sc. Nat., ser. 6, xiii (1882), page 9. 3o6 VEGETATION OF THE NEW JERSEY PINE-BARRENS leaves of the laurel are elliptic and leathery in the sun-plant. The leaves from a bush growing in the shade are larger, thinner and longer. The pyxie, Pyxidanthera barbulata, has a sun form and a shade form. Figure 279 Influence of sun and shade on pine-barren plants A, Leaf of laurel, Kalmia latifolia, from plant growing in bright sun and dry soil. B, Leaf of laurel, Kalmia latifolia, from plant growing in the shade. C, Bidens trichosperma growing in deep shade in Cedar Bog, two miles west of Pt. Pleas- ant, N. J., September 9, 1912. D, Bidens trichosperma growing in medium shade in Cedar Bog. E, Bidens trichosperma growing in bright sunny places. F, Py.xie, Pyxidanthera barbulata, growing in bright sun and dry sandy soil. G, Pyxie, Pyxidanthera barbulata, growing in dense shade and leaf mould. VEGETATION OF THE NEW JERSEY PINE-BARRENS 307 The former is denser and more cushion-like with smaller leaves, the latter is more open, less compact in growth with larger leaves (Fig. 279). (c) Wind The wind-shearing action on trees is noticeable, especially in the pitch- pine trees, Pinus rigida. In the coastal pine-barrens, the action of the wind in producing one-sided trees and partially prostrate trees is worthy of note. The action of the wind on the vegetation of the plains has been described. (Figs. 31, 32, 33, 34, 35, 36, 37.) (d) Water Quite a few pine-barren plants have forms that are due to the action of water. Most plants of Sagittaria graminea have leaves that are lanceolate, but in wholly submerged plants the leaf-blades are partly, or entirely absent, being represented by linear phyl- lodia. The water club-rush, Scirpus subterminalis, has in its usual submerged forms long, narrow leaves, streaming in the direction of the current of water. In ponds where the water has been drained off it often grows upright, with much shorter and stiffer stems. Eriocaulon sep- tangulare, in its submerged form, has well-developed leaves, about as long as the scape. Other leaves are only half the length of the scape, while plants on the edge of a pond or swamp are often 75 to 100 millimeters in height, with leaves 25 millimeters long, according to Stone's statement (324). The bayonet-rush, J uncus militaris, has curious, submerged, thread- like leaves that arise from the root-stock and grow in the water-like masses of waving hair. The low water-milfoil, Myriophyllum humile. Figure 280 Leaf forms of several species of pond weeds, Potamogeton. ard Long at Man- ahawkin, Septem- ber I, 1907. Potamogeton Oake- sianus with broad floating leaves, collected by Bay- ard Long at Fol- som along Hospi- tality branch of Great Egg Har- bor River, July 27, 1909. A, Potamogeton con- fervoides with narrow linear sub- merged leaves, collected by Bay- ard Long at Bam- ber, May 8, 1907. B, Potamogeton epi- hydrus with (a) narrow linear sub- merged leaves and (b) broad floating leaves; collected by Bay- 3o8 VEGETATION OF THE NEW JERSEY PINE-BARRENS exists in the form of small plants creeping in the mud (humile) or in a submerged form (capillacea), and those with an emersed spike (natans). Several species of Potamogeton and Utricularia show the influence of an aquatic environment on the form and structure of the plant body (Figs. 280 and 281). Convergent Epharmony. — It is evident that many growth forms among the pine-barren plants may be referred with confidence to the influence of the stimuli of various external factors. Some of these forms are mere fluctuations, or oscillations, about a norm, others are hered- itary and remain constant. Cockayne in his study of New Zealand vegetation enunciates a principle applicable also to the New Jersey pine- barren vegetation, that, "It is a fact of the greatest significance that iden- tical forms are found side by side amongst species belonging to unrelated families." A few illustrations of this principle will be mentioned, as they have come within the notice of the writer, as cases of convergent ephar- mony. (a) Annual Wiry Stems, Repeatedly Branched Anychia canadensis is an annual herb of the family Caryophyllaceae. Crotonopsis linearis is an annual herb of the family Euphorbiaceae. In growth forms they are much alike. The first plant has a very slender, filiform, usually erect stem, which is forked repeatedly with opposite, small, elliptic, oval, or sometimes oblanceolate leaves. The second plant has wiry stems, repeatedly forked, and bearing small oblong-ovate to linear-lanceolate leaves. It can be distinguished in the vegetative con- dition from the first by its silvery-scurfy appearance. Both plants are of about an equal height. (h) Annual Herbs with Articulate Stem and Subulate Leaves Three herbs approach each other in growth forms. They are Poly- gonella articulata (Polygonaceae), Sarothra gentianoides (Hypericaceae) and Bartonia virginica (Gentianaceae). The first plant is an annual herb with slender, jointed, wiry, erect stem diffusely spreading, or branched with linear or linear-subulate leaves. The plant is found in barren, sandy places in the coastal pine-barrens. Sarothra gentian- oides is an erect annual fastigiately branched, and with wiry branches. The leaves are minute and subulate. It also grows in sandy soil, Bar- tonia virginica is an erect, stiff annual with filiform, wiry stem beset with leaves that are reduced to subulate scales appressed to the stem. It is found in moist, sandy soil. VEGETATION OF THE NEW JERSEY PINE-BARRENS 309 Figure 281 Details of submerged and aerial parts of several species of Utricularia. Drawings from specimens in the Herbarium of the Philadelphia Botanical Club, where data will be found. A, Utricularia clandestina. B, U. cleistogama. C, U. cornuta. D, U. inflata. E, U. juncea. F, U. subulata. G, U. intermedia. H, U. fibrosa. I, U. purpurea. 310 VEGETATION OF THE NEW JERSEY PINE-BARRENS (c) Perennial Heath-like Plants The pine-barren sand-wort, Arenaria caroliniana (Caryophyllaceae), and the heath-like Hudsonia ericoides (Cistaceae), are convergent forms (Fig. 282). The first is a perennial herb from a deep root and with tufted, erect stems covered with rigid, subulate leaves. Hudsonia eri- coides is bushy-branched from the base. Its leaves are subulate. Both forms are suggestive of each other, especially the young plants. (d) Hassock-like Growth Forms The idea of a hassock is a thick, rounded cushion, used as a foot-stool. It implies a besom, or bushy object, while a cushion growth form in the ecologic sense is like a pin-cushion, as we find in such plants as Pyxi- danthera barbulata of the pine- barrens, or Silene acaulis of al- pine regions. The growth form now to be described may be compared to a globular bush, which owes its form to the constant clipping by shears in the hands of a gar- dener. This form in nature is induced in a dry soil with strong transpiration, so that the plant has short curved, or crooked shoots and stems with short internodes and interlaced branches. That dryness induces these growth forms is proved by the fact that abundant moisture, when the growth habit is not fixed, causes an elongation of the shoots with a lengthening of the internodes. Two pine-barren shrubs illustrate this form to perfection, viz., Corema Con- radii (Empetraceae) and Dendrium (Leiophyllum) buxifolium (Erica- ceae). The first shrub native to the plains exists in great round masses, i to 3 feet in diameter, the basal portion a tangle of brown stems and dead branches, but the surface covered with the little, slender, needle-like leaves. The sand-myrtle, Dendrium buxifolium, is a low, evergreen shrub resembling a low box-bush, but its leaves are larger, although tough and leathery. (e) Prostrate Shoots Many species growing in dry, warm, sandy soil and in other habitats have prostrate shoots. A few examples from different families may be Figure 282 Convergent Epharmony in Sandwort, Aren aria caroliniana and Hudsonia ericoides. VEGETATION OF THE NEW JERSEY PINE-BARRENS 3II cited from the pine-barren region of New Jersey. The following are of interest: Epigaea repens (Fig. 193), Gaultheria procumbens (Fig. 188), Arctostaphylos uva-ursi (Ericaceae), (Fig. 197), Euphorbia ipecac- uanhae (Euphorbiaceae), Galactia regularis (Leguminosae) (Fig. 180). (/) LlANES The climbing plants of the New Jersey pine-barrens belong prominently to five genera of four families of flowering plants. In the climbing habit they approach each other, although of diverse morphologic structure. They are species of Smilax (Smilacaceae), Dioscorea villosa (Dioscorea- ceae), Vitis aestivalis, Ampelopsis (Psedera) quinquefolia (Vitaceae), and Rhus radicans (Anacardiaceae). Species of Ipomoea, Convolvu- lus, Cuscuta (Convolvulaceae), have twining stems, while two legu- minous plants, Clitoria mariana * with twining habit, and Apios tuberosa, with climbing habit, are herbs to be included in this category. (g) Plants with Aerenchyma Five aquatic plants, belonging to four different families of flowering plants, possess aerenchyma. They are Hypericum adpressum (Hyper- icaceae) found at May's Landing at the edge of the pine-barrens, Decodon verticillatus (Lythraceae), Rhexia virginica and R. aristosa (Melastomaceae) and Ludvigia sphaerocarpa (Onagraceae). Aeren- chyma is a tissue-like cork, which has its own phellogen, but it consists of thin-walled, non-suberized cells between which are large air-contain- ing, intercellular spaces. The tissue shows itself as a white, spongy covering. In the five plants above mentioned, it is found around the bases of the stems and as in a number of other helophytes, or swamp plants in other parts of the world. (h) Cleistogamy A number of plants native to the pine-barren region are characterized by the formation of underground cleistogamous flowers which produce normally a larger number of good seeds than the conspicuous flowers in an aerial position. Such cleistogenes are Amphicarpon Purshii (Graminaceae), Viola sagittata, V. emarginata, V. primulifolia, V. lanceolata (Violaceae) (Fig. 283) and Utricularia cleistogama (Lenti- bulariaceae) (Fig. 281 B). In the cleistogamous flowers of the violets, see figures (Fig. 283) * Very frequently erect, not twining. 312 VEGETATION OF THE NEW JERSEY PINE-BARRENS adapted from Church,* the pollen grains are fewer in number than in normal flowers and the ovules are about half the normal number. The anthers lie against the ovary wall, and the distance from the uppermost pollen grains to the end of the rudimentary style is extremely small (about 5 or 6 diameters of the pollen grain). The pollen grains germi- nate within the pollen sac and the wall of the anther box is ruptured and the pollen tubes grow across to the style which they penetrate, thus accomplishing the act of close pollination. The list of converging growth forms which have been described in categories (a) to {h) inclusive is not exhaustive, because we have omitted a description of a number of distinct kinds, such as the rosette form. The above account, however, will serve as an introduction to a very attractive field of ecologic investi- gation. Persistent Juvenile Forms Cockayne t states that about 200 species of New Zealand vascular plants belonging to 37 families show a more or less well-marked distinction between the juvenile and adult stages of development while, perhaps, in 100 species the differences are very great indeed. In the absence of exact data for the pine-barrens of New Jersey, and to draw attention to this important phase of evolutionary investi- gation, a single example of the persistence of a juvenile form may be cited. The red cedar, Juniperus virginiana, in the early stages of growth has awl-shaped, divergent leaves, which in most trees are replaced by closely appressed scale leaves. In a few trees which 1 have seen in the coastal plain of New Jersey the juvenile state persists until the tree reaches a considerable size. Herbarium specimens of these persistent juvenile forms have been preserved. One way in which these persistent juvenile forms differ from the adult forms is in the bright green color of the foliage, as contrasted with the darker greens of the tree, with closely appressed scale leaves (Fig. 284). * Church, A. H.: Types of Floral Mechanism, Part 1, page 98 (1908). t Cockayne, L.: Observations concerning Evolution derived from Ecological Studies in New Zealand. Trans. New Zealand Institute, xliv, 191 1. Figure 283 Cleistogamous flowers of violet. Ovules. Growth of pollen from anther across to stigma (Cleisto- gamous pollina- a, Inturned sepals. b, Covering inturned petals. c, Stamen and anther with pollen. d, Stigmatic region. f. tion). VEGETATION OF THE NEW JERSEY PINE-BARRENS 313 Hybridization A number of natural hybrids have been collected in the pine-barren region of New Jersey. The white fringed orchid, Blephariglottis blephariglot- tis, is concerned as one of the parents in the formation of two hybrids. Blephariglottis bicolor Rafmesque is B. blephariglottis X B. ciliaris, and has been found at Bamber by Bayard Long on August 25, 1909, according to Witmer Stone. B. Canbyi Ames is a hybrid of B. blephariglottis and B. cristata, and has been collected a number of times in the pine-barrens. A hybrid between Quercus phellos and Q. ilicifolia was found by J. E. Peters at May's Land- ing, while east of Farmingdale, at the northern edge of the pine-bar- ren region, the writer found Quer- cus heterophylla, the Bartram oak, which has been proved by the experiments of MacDougal * to be a hybrid of Quercus phellos and Quercus rubra as parental forms. The fifth case of a natural hy- brid that has come to my notice is one produced in a natural cross between Eupatorium resinosum and E. perfoliatum discovered by a former student of mine, Bayard Long of the Philadelphia Botanical Club. Figure 2 Juvenile and adult leaf forms in pine and juniper. A, Winged seed of Pi- F nus rigida. B, Winged seed of Pi- nus echinata. C, Seed of white cedar Chamaecyparis thyoides. D, Piece of a sprout from a fire-injured pitch pine, show- ing (a) reverted ju- venile leaves; (b) sheathing leaves; (c) secondary nee- dle leaves. Adult shoot of red cedar, Juniperus virginiana with berries. Young shoot of red cedar, Juniperus virginiana, show- ing persistence of juvenile need le- like leaves. Conclusion The above evolutionary studies have been treated in a general way and from the observational standpoint. The field has not been ex- hausted and much important work remains to be done in the field * IVIacDougal, D. T.: Hybridization of Wild Plants. The Botanical Gazette, 43: 53, January, 1907. 314 VEGETATION OF THE NEW JERSEY PINE-BARRENS under experimental conditions. The above account is, therefore, a short introduction to a subject of considerable botanic interest. It provides an outline of what may be accomplished by intensive methods of research work, spread over a period of years. Preferably the investi- gation should be conducted in the field under controlled conditions, rather than in the laboratory where even with the best surroundings the environmental factors are purely artificial. An ideal system would be to have plots of land, an hectare or two in extent, located in the pine forest, in the plains, in a cedar swamp, and in a deciduous swamp, sur- rounded by a barbed-wire fence 2 to 3 meters in height, so as to protect the experimental ground. Within the inclosures the instruments de- voted to the investigation of the climatic and other factors could be set up with a reasonable hope of their remaining undisturbed during the course of the research work. The foregoing survey of the vegetation of the pine-barrens of New Jersey, although not exhaustive, presents some interesting facts about a vegetation which, although related to that of the southern coastal plain, yet has many features that are entirely distinct. We have seen in the first place that in no other region of North America does the pitch pine, Pinus rigida, cover such an extended area of country as a dominant tree. The pine-barrens of the southern states, as we have seen, are characterized by the dominance of other pine species, notably the long leaf-pine, Pinus palustris, the loblolly-pine, Pinus taeda, the yellow- pine, Pinus echinata, and the slash-pine, Pinus caribaea. These are all dominant in the area of country representing their natural range, and on soils which are different for each of the pine species above men- tioned. Secondly, the pine-barren vegetation of New Jersey is an old vegeta- tion in a long occupancy of the territory in which it is found. The age of this forest is attested by its survival through the changes of several geologic epochs antedating the glacial period, when the northern part of the State was covered with ice. Thirdly, the pine-barren vegetation has maintained itself against the encroachment of other types of vegetation. This is attested by the difficulty that introduced plants have in gaining headway against the ex- clusive closed formation of pine-barren plants, and even today, after long occupancy of the region by man, introduced weeds play a relatively unimportant part in the flora. Fourthly, nowhere in the northern United States, east of the moun- tains, is there such a large geographic area which exists in such a wild and pristine state. This is due undoubtedly to the rapid settlement of VEGETATION OF THE NEW JERSEY PINE-BARRENS 315 the richer and more desirable lands of the western states, which have yielded larger financial returns with a smaller outlay of capital. The pine-barren soils, although suitable for the growth of many vegetable crops, yet yield their returns only after an expenditure of money and labor which until recent years has not been considered commensurate with the returns. With the settlement of the western states, attention will be turned to the poorer sandy soils of these northern pine-barrens, which yield fair returns if properly treated. Already the effect of the settlement of Italians and Russian Jews in the southern part of the terri- tory is felt in the removal of the pine forests and in the cultivation of the soil. As early as 1888 the following statement was made: "The remainder of the region is being rapidly improved and brought under cultivation. Twenty-five per cent, of the pine region is cleared in Camden County, 30 per cent, in Gloucester, and 63 per cent, in Salem. Perhaps the most advancing portion of the region is Cumber- land County, where 36 per cent, is now cleared. It is true that a good part of this is the older settled country west of the Cohansey, but large inroads have been made in the eastern portion within 25 years, particu- larly in the vicinity of Vineland." To this statement might be added one, that at Woodbine, large areas have been cleared by the Baron de Hirsch estate, and rapid inroads have been made in the pine forest along the railroad between Newfield and May's Landing. Fifthly, the future development of the region, according to the writer's acquaintance with it will be along four lines. Agriculture and horti- culture will remain always the most important industries of the pine region. The preservation of the pine forest in large tracts will be neces- sary to preserve a proper balance between forested areas and agricul- tural areas. At least 30 per cent, of the region should be kept perma- nently in the form of state forests, owned and managed by the State in perpetuity. Manufacturing interests will be centered in the larger towns, where the factory employee will be within reach of his own home ground, where supplementary supplies of food can be raised in his own kitchen-garden. Lastly, the region, having a salubrious climate, will naturally attract the health seeker and sufficient stretches of pine-forest should be pre- served for the benefit of this class of the community. The future de- velopment of the region into one of unusual prosperity is only a matter of time. Its close proximity to the vast populations that center about New York City and Philadelphia, will make ultimately of the region a valuable agricultural country. The State of New Jersey has begun to have a far-sighted policy in the development of its roads and its inland 3l6 VEGETATION OF THE NEW JERSEY PINE-BARRENS waterways. In addition to these improvements the State should take over large blocks of forest for the future use and pleasure of its people. 1 ts streams should be kept from pollution and stocked with fish. The larger forest tracts should be stocked with game and forest paths con- structed, so that the largest number of people, healthy and sick, wealthy and poor, could derive benefit from the life-giving air of the pines. NDEX ^n almost complete list of pine-barren plants, 548 species out of 555 native ones, will be ifound on pages 196-208. As these names are arranged systematically, they are not included in the index. Additional lists of plants will be found on pages 83, 127, 139, 145, 172, 180, 186, 189, 190. Abama americana, roots and stems of, 225 Abbe, Cleveland, cited, 194 Acer carolinianum, 305; figure of micro- scopic leaf structure, 274, micro- scopic leaf structure, 274 rubrum, 112, 133, 134, 137. '4o; mu- tation of, 305 tridens, 303 Acorn vivipary, 292 Additional pond plants, 144 swamp plants, 138 Adier, H., cited, 293 Aerenchyma, 31 1 Age of larger pines of Lower Plain, 133 of pine sprouts on plains, 153 of tree sprouts of plains, 134 Agriculture, future, 313 Aletris farinosa, forms of, 303 Algoid leaves, 233 Allen, Grant, cited, 188 Alnus rugosa, figure of microscopic leaf section, 268; microscopic leaf structure, 268 Alpine meadows, study of, 173 Amentiferous trees, flowering period of, 21 1 American and European plant formations compared, 168 Ampelopsis quinquefolia as liane, 31 1 Amphibolips coelebs, the oak spindle gall, 297 confluentus, gall insect on scarlet oak, 296 Amphicarpon Purshii as a cleistogene, 3 1 1 Analyses of white-cedar trees, 1 19 Andricus seminator, gall on white oak, 296, Andromeda ligustrina, figure of microscopic leaf structure, 280; micro- scopic leaf structure, 280 mariana, figures of, 237; root system of, 237; microscopic leaf structure, 280 Andropogon scoparius, roots and stems of, 224, 223 Animals of pine-barrens, 13 Annual herbs in marshes, 191; of pine- barren formation, 187 Anthropochores, 106 Anthropophile element, 106 Anthropophytes, 106 Apios tuberosa with twining habit, 3 1 1 Apocynum medium as a species, 300 Milleri as a species, 300 urceolifer as a species, 300 Apophytes, 106 Apparatus used in soil research, 36 Application of fertilizer to soils of plains, 163 Aquatic plants in cedar swamps, 190 Arbutin as a reserve food, 276 Archaeophytes, 107 Architectural form of pine trees, 53 Arctostaphylos heath, 168 Arctostaphylos uva-ursi, 63, 68, 3 11 ; figure of, 241; figure of microscopic leaf struc- ture, 276; microscopic leaf structure, 273 ; photograph of, 1 57; root system of, 239 Area of pine-barrens, 178 Arenaria caroliniana, 85; figure of, 310; figure of microscopic leaf struc- ture, 270; microscopic leaf structure, 270; heath-like form, 310 peploides var. robusta, 5 Artificial ponds, 144 Asclepias amplexicaulis, figure of, 234; root system of, 234 obtusifolia, root system of, 234 Aspects of vegetation, 180 seasonal, of white-cedar swamps, 127 Association coefficient, 175 Associations in cedar swamp stream, 123 Aster nemoralis, 3 radula, 3 Atlantic Highlands, 3 .Autumn aspects, 80; of cedar swamps, 128 leaf colors, 195, 209 Average rate of growth of white-cedar, 119 Avivectent seeds, 93 Azalea viscosa, 112, 134, 133, 137; figure of microscopic leaf structure, 276; micro- scopic leaf structure, 276 317 3i8 VEGETATION OF THE NEW JERSEY PINE-BARRENS Baccharis halimifolia, 95 Bailey, L. H., book by, 252 Balsams, 28 Baptisia tinctoria, figure of, 233; root sys- tem of, 233 Bark for drugs, 28 of pitch-pine, 98 Barren grounds of Canada, 9 Barren Hill, 9 Barrens, application of term, 9 Bartonia virginica, 308 Basket form of pitch-pines, 1 52 Bassett, William F., cited, 172 Bastard toad-flax, root system of, 230 Bayberry, microscopic leaf structure, 268 root system of, 228 Bay-like leaves, 235 Beacon Hill area, 3; formation, i Bearberry, 65, 68; microscopic leaf struc- ture, 275; photograph of, 157; root sys- tem of, 239 Bear-oak, microscopic leaf structure, 268; root system of, 230 Beauchamp, William M., cited, 10 Bermuda dune plants, 285 Beutenmiiller, William, cited, 295, 296 Beyerinck, M. W., quoted as to galls, 295 Bidens trichosperma, sun and shade forms, 306 Biologic spectrum, 186 types, 186; of marsh formations, 190; of pine-barren plants, 185 Bird-distributed seeds, 93 Bird-foot violet, mutations, 304 Birkenheide, 86, 89 Black-jack oak, 89, loi; microscopic leaf structure, 269; root system of, 230 Blackman, Leah, quoted, 14 Black oak, 102 Bladder leaves, 257 Bladderwort, figures of, 309 Blakeslee, Albert F., cited, 53 Blephariglottisblephariglottis X B.ciliaris, 313 blephariglottis X B. cris- tata, 313 Canbyi, a hybrid, 313 Block Island, 5 Blueberry, 85, 133, 134; culture, 238; root system of, 238 Bluff forest at Somers Point, 91 Bog asphodel, roots and stems of, 225 Bog formation, 48 Bog plants, additional list of, 126 Bogs, 26 of northern New Jersey, 1 1 5 Boulder plants, 219 Boxing pine trees for turpentine, 22, 23 Bracken, 133; roots and stems of, 220 Branch swamps, 136 Briggs, L. J., cited, 45, 47 Broom-crowberry, 151, 304; microscopic leaf structure, 273 Brown, Harry P., cited, 209 Stewardson, cited, 156 Bulbous shoots, 218 Bush fruits, 173 Caatingas, 145 Caleche of Mexico, 166 Calico-bush-, mutations, 304 Calopogon pulchellus, roots and stems of, 227 Calvert, P. P., cited, 296 Campos of Brazil, 145 Cannon, W. A., cited, 217, 220; his classi- fication of roots, 247 Canopy of pine forest, 60 Cape Cod, 5; plants of, 5 Cape May deposits, 5 Capillary rise of water, 42 Carex hormathodes, 5 silicea, 5 sterilis, 5 trisperma var. Billingsii, 5 Carolinian types, 5 Cassandra calyculata, 143; in savanna, 149; root system of, 239 Cassia chamaecrista, root system of, 233 Castalia odorata, 144; figure of microscopic leaf structure, 271; figure of microscopic petiole structure, 271; flowering and fruiting of, 210; microscopic leaf struc- ture, 270; mutations, 304; photograph of, 123; root system of, 232 Caudex multiceps, 218 Caudices, 217 Cause of dwarf trees of Coremal, 159 of galls, 295 Cecidology, 293 Cecidomyia pisum, 297 Cedar (red) leaf forms, 253 Cedar swamp formation, 48, 114, 189; blending with deciduous swamp, 134 Cedar swamp at Lakehurst, 1 14; at Mays Landing, illustration of, i 16; between Whitings and Bamber, illustration of, 121; blending with pine forest, 112; synecology of, 120; yield of, 20 Census of plants in lots at Belmar, 83 of vegetation, 173 Cereal crops, 173 Chamaecyparis thyoides, 112; base of, 118; figure of microscopic section of leaf, 264; microscopic leaf structure, 264; photo- graph of, 126 Chamaedaphne calyculata, figure of micro- scopic leaf structure, 277; microscopic leaf structure, 277; root system of, 239 Chamaephytes, 185 Chamberlain, T. C, cited, i VEGETATION OF THE NEW JERSEY PINE-BARRENS 319 Charcoal, 20 Chemic analysis of soils, 34 Chimaphila maculata, figure of, 235; root system of, 236 Chrysopsis falcata, figure of microscopic leaf structure, 282; micro- scopfc leaf structure, 282 mariana, figure of, 243; figure of microscopic leaf structure, 283; microscopic leaf struc- ture, 283; root system of, 242 Church, A. H., cited, 312 Cinnamon-fern, 80, 134, 133 Circinate ptyxis, 253 Cladonia rangiferina, 132 Clammy azalea, 134 Clasping leaves, 257 Classification of alien plants, 106; of leaf structures, 260; by root systems, 246; of root systems by Cannon, 220; of shoots by Hess, 217, 218, 219 Cleistogamy, 311; in violet, figures of, 312 Clements, F. E., cited, 175, 259 Clethra alnifolia, 112, 137; figure of, 233; figure of microscopic leaf structure, 277; microscopic leaf structure, 277; root sys- tem of, 235 Climbing plants in marshes, 191 Clitoria mariana, with twining stems, 3 1 1 Coastal oak forest, 104 pine-barrens, 82 Cockayne, L., cited, 298, 308, 312 Coefficient of association, 173; of genera, 176, 181 Cole, Rex Vicat, cited, 33 Collection of turpentine, 22 Colors of flowers in white-cedar swamps, 190; of marsh plants, 92; tabu- lated, 188 Colors of leaves in autumn, 195, 209 Colors of pine-barren vegetation, 77 Columbia formation, i Comandra umbellata, root system of, 230 Commencement of cambial activity in pitch-pine, 209, 210 Comparison of American and European plant formations, 168 of quadrats, 180 Competition of root systems, 231 Compound palmate leaves, 236; pinnate leaves, 256; trifoliate leaves, 256 Comptonia asplenifolia, 67, 83; figure of, 228; figures of leaf sections, 268; micro- scopic leaf structure, 267; root system of, 228 Conard, Henry S., cited, 210 Conclusion, 313 Conditions in cedar swamps, 1 15 Cone production, law of, 292; on different trees, 287; of pitch-pine, 286 Cones of pitch-pine, figure of, 286 Connold, Edward T., cited, 293 Convergent epharmony, 308 Convolvulus with twining stem, 3 1 1 Cook, Mel T., cited, 295 Corema Conradii, 3, 6, 143, 151, 133, 169; botanic characters of, 136; figure of mi- croscopic leaf structure, 274; forms of, 304; growth formS, 310; microscopic leaf structure, 273; photograph of, 132, 157, 138; on Nantucket, 155; time of flowering, 1 56 Coremal, 151, 138, 169; cause of dwarf trees, 159; measurement of species, 139; of New Jersey, 168; on Nantucket, 165; size of plants, 139; soils treated experi- mentally, 162; synecology of, 134; syn- opsis concerning, 168; theories concern- ing, 159 Corn culture in soil from plains, 162, 163, 163; in top-soil, 169 Coville, F. v., cited, 238 Cowbane as an elementary species, 299 Cow-lily, root system of, 231 Cow-wheat, root system of, 242 Cranberry, 23 bog, 25; formation, 48 culture, 25 sorting house, 26 storage house, 26 Cranmer, George, cited, 160 Crested shoots, 218 Crotonopsis linearis, 308 Cryptophytes, 185 Crystals, rhomboidal, 262 Cultivated plants of the pine-barrens, 172 trees of the pine-barrens, 172 vines, 173 Culture of corn in plains soils, photograph of, 162, 163, 165; in top-soil, 169 of cranberry, 23 of peas in soil from plains, 162; photograph of, 163; in top-soil, 169 of sunflowers in soils from plains, 164, 167 of wheat in soils from plains, 164 Cup-and-gutter method of turpentine gath- ering, 22 Curves showing percolation of water, 39 Cuscuta with twining stems, 31 1 Cuticle thick in leaves, 260 thin in leaves, 260 Cypripedium acaule, roots and stems of, 226, 227 Dams, 144 Dandelion leaf form, 255 Darwin's discussion of variation, 300 Date of pollination of pitch-pine, 209 Davis, Charles A., cited, 1 17 320 VEGETATION OF THE NEW JERSEY PINE-BARRENS Deciduous-leaved species, i88 leaves, 255, 262 swamp at Belmar, photograph of, 135; swamp and cedar swamp blending, 134; swamp formation, 48, 133; photo- graph of, 132; transition to, 132; of transition region, 136 trees in cedar swamps, 189; in marshes, 192 vegetation of Delaware Valley, 9 Decodon verticillatus with aerenchyma, 3 1 1 Defebaugh, J. E., cited, 19 Delgar, Colonel E., of Denmark, work of, 161 Demarcation between plains and pine for- est, 158 Dendrium buxifolium, 7 1, 72; figure of, 236; figure of microscopic leaf structure, 280; growth forms, 310; microscopic leaf structure, 280; root system of, 236 Denmark, reforestation of, 161 Density, floral, 176 Detailed root and stem study, 219 et seq. Development of pitch-pine seeds, 209 De Vries, Hugo, cited, 298, 299; views, 300 Diageic plants, 185 Diagram of savanna, 147; showing succes- sion, 49 Diameter of pitch-pine trees, 288, 290, 291 Dioscorea villosa as liane, 3 1 1 Diphotic leaves, 259 Diphotophyll, 259, 262 Diplophyll, 259, 263 Diseases of white cedar, 129 Dispersal of coastal plain plants, 6 Dissected submerged leaves, 256 Distribution of roots, 245 of transition plants, 100 Districts, floral, 15 Dodder leaf form, 257 Dormancy of plants, 195 Drosera rotundifolia, figure of leaf tentacle, 272; microscopic leaf structure, 272 Drought, relation of plants to, 249 Drude, Dr. O., cited, 190 Drug plants, 27, 28 Dulichium spathaceum, roots and stems of, 224 Dumb-watch, 17 Dwarf chestnut-oak, root system of, 230 pine trees of plains, 167 trees of Coremal, cause of, 159; of Hempstead Plains, 171 Early glacial time, 6 saw mills, 18 settlements in West Jersey, 13 East Plains, 15 i Edinger's drawing apparatus, 263 Eichenheide, 86, 168 Elatine americana, 5 Elementary species, 298, 299 Elevations in plains, 151 Engler, Prof. Adolf, 63 Enumeration of pine-barren plant families, 182, 183 Epharmony, 305; convergent, 308 Ephemerophytes, 107 Epidermis, papillate, 261; thick-walled, in leaves, 260 Epigaea repens, 311; figure of, 239; figures of microscopic leaf structure, 277; micro- scopic leaf structure, 277; root systems of, 237 Epigeic plants, 185 Epokophytes, 107 Ergasiolipophytes, 106 Ergasiophygophytes, 107 Ergasiophytes, 106 Eriocaulon septangulare, 5; floating and submerged forms, 307 Eriophorum virginicum in savanna, 148 Eupatorium resinosum X E. perfoliatum, 3'3 verbenaefolium, microscopic leaf structure, 283; root system of, 242, 244 Euphorbia ipecacuanhae, figure of, 234, 299; figure of microscopic leaf structure, 273; as an elementary species, 299; micro- scopic leaf structure, 273 ; root system of, 234 European and American plant formations compared, 168 Evergreen leathery leaves, 236 leaves, 262 species of pine-barrens, 188 trees in cedar swamps, 189; in marshes, 191 Evolutionary viewpoint, 298 Exotic place names, 1 1 Experimental inclosures, 314 treatment of Coremal soils, 162 Experiments with soils, 35 Facies of pine-barren formation, 56; of pine-barren vegetation, 57 Factors influencing regeneration of pitch- pine, 291, 292 Fall of pine-barren streams, 8 False asphodel, roots and stems of, 225 Families of pine-barren plants, 182, 183 Fascicled leaves, 253 Fernald, M. L., cited, 5 Ferns in cedar swamps, 189; in marsh for- mations, 190; of pine-barren formation, 187 Fertility of soils, theories, 32 Fertilization of egg-cell in pitch-pine, 209 of pine, 291 VEGETATION OF THE NEW JERSEY PINE-BARRENS 321 Fertilizer, application to plains soils, 165 Field controlled experiments, 314 Fire-denuded pitch-pines, 51, 52 Fire, influence in formation of plains, 169 strips, 100 Fires on plains, 1 59 Flat pine-barren facies, 63 Floating leaves, 256 Floral density, 176; districts, 15; proces- sion, 193 Flower colors, numbers of species with, 189; of lupine, 303; of marsh plants, 192; summarized, 192; tabu- lated, 188; of white-cedar swamp plants, 190 crops, 173 Flowering and fruiting of Castalia odorata, 210; of pine-barren plants, 210; of amentiferous trees, 211; of Pyxidanthera barbulata, 210 moss, 152; root system of, 240 periods, 193; of pine-barren plants, 196 et seq. Flowers, 29; numbers of different months, 212, 213; procession of, 213, 214, 215 Fluctuating variations, 300 Fluctuations of plants, 300 Fly-trap leaves, 237 Fodder crops, 173 Forest canopy, 60 Formation of bogs, 117; of plains, 151; of pond plants, 143; of savannas, 143 Formations of America and Europe com- pared, 168; phytogeographic, 48 Forms of leaves, 252 Frequency of individuals, 176 Fritsch, F. E., cited, 216 Frost, first killing, 194; last killing, 194 Fruit of Pyxidanthera barbulata, 210 Fruit trees of pine-barrens, 172 Fruiting period of plants, 193, 196 et seq. Fruits, cultivated bush, 173; small, 173 Fungi on white-cedar, 131 Fungous diseases of white-cedar, 129, 130, '3" Future of pine-barren region, 3 i 5 Galactia regularis, 311; figure of, 233; root system of, 233 Gall-gnats, 297 Gall-midges, 297 Galls, cause of, 295; figures of, 296; of insects, 293 Game preserve, 316 Garden crops, 174 Gates, Frank C., cited, 239 Gathering of flowers, 29; of greens, 29 Gaultheria procumbens, 311; figure of, 237; figure of microscopic leaf structure, 278; microscopic leaf structure, 278; root sys- tem of, 236 ~^ 21 Gaylussacia baccata ( = resinosa), micro- scopic leaf structure, 279 dumosa, figure of microscopic leaf structure, 278; micro- scopic structure of leaf, 278 dumosa var. Bigelowiana, 5 frondosa, 69, 112, 135; figures of microscopic leaf structures, 278; microscopic leaf struc- ture, 278 resinosa, 69, 73, loi; figure of, 240; figure of microscopic leaf structure, 279; micro- scopic leaf structure, 279; root system of, 238 General observations on pine-barren vege- tation, 173 remarks on leaf forms, 258 Generic coefficient, 176, 181 Geographic location of pine-barrens, 8; of plains, 169 place names, 9 Geography of pine-barren region, 8 Geologic map of pine-barren region, 3 Geophytes, 185 German plant formations, 85 Germination of unripe acorns, 293 Gifford, John, cited, 1 1 Glabrous leaves, 261 Glacial period, 1 Gnarled trees of plains, 1 56 Goat's-rue, root system of, 232 Goebel, K., cited, 252 Golden aster, root system of, 242 Golden-club, 147; leaf structure, 263 Graebner, Dr. Paul, cited, 48, 83, 166, 167 Grand period of growth, 194 Graph of leaf structure, 283; representing water-retentiveness, 44; showing rate of percolation, 39 Grass form of leaves, 253 Grasses in cedar swamps, 189; in marsh formations, 190 Great Egg Harbor River, 7 Greens, 29 Gridiron method, 176 Griggs, Robert F., cited, 289 Groom, Percy, cited, 249 Growing season, 194 Growth in pitch-pine stem, 210 of white cedars, average rate of, 119 Guard cells deeply depressed, 261; project- ing, 261; present at surface, 261; slightly depressed, 261 Guppy, H. B., cited, 292, 293 Haas, Paul, and Hill, T. G., cited, 276 Habenaria blephariglottis, 3; roots and stems of, 227 Haberlandt, Prof. G., cited, 53 Hall, A. D., cited, 32 322 VEGETATION OF THE NEW JERSEY PINE-BARRENS Hall, William L., cited, 19 Hapaxanthic species, 217 Harper, Roland M., cited, 108, 144, 170, 182; photograph by, 140 Harshberger, J. W., cited, 4, 9, 15, 17, 25, 48, 129, 161, 181, 259 Hassock-like growths, 310 Hawley, L. F., cited, 23 Heat of soils, 250 Heather leaf forms, 253 Heathland, 166; of Nantucket, 168, 170; photograph of, in Nantucket, 170; of New Jersey, 170; physiognomy of, 167 Heath-like plants, 310 Helophytes, 185 Hemicryptophytes, 185 Hempstead Plains, 108, 170 Henkel, Alice, cited, 29 Herbs, annual, 187; as drugs, 28; with articulate stem, 308; perennial, 187; with subulate leaves, 308 Herty, Charles H., cited, 22 Hess, Eugen, cited, 217 Hieracium venosum, figure of, 242; root system of, 242 High pine-barren facies, 60 Hillman, Sarah C, cited, 25 Holcaspis centricola on Quercus stellata, 297 Holly, 90, 91 Holm, Theo., cited, 226 Hopkins, Albert A., cited, 24 Horticulture, future, 315 Huckleberry picking, 27; root system of, 238 Hudsonia ericoides, 5, 75, 112, 310; figure of, 231, 310; figures of micro- scopic leaf structure, 273; mi- croscopic leaf structure, 275; root system of, 23 1 tomentosa, figure of, 231; root system of, 230 Huntington, Annie Oakes, cited, 53 Hybridization, 3 13 Hydrophytes, 185 Hypericum adpressum with aerenchyma, 3" densiflorum, 140; figure of, 232; root system of, 23 i Hypodermis in leaves, 260 1 DioBLASTS, 262 ; in leaves of water-lily, 27 1 Ilex glabra, 69, 133; figures of microscopic leaf structure, 274; microscopic leaf structure, 274; root system of, 234 opaca, 90, 91, 94, 95, 96 Impervious subsoil of the plains, 169 Importance of plant families, 183, 184 Inaccessibility of pine-barrens, 12 Indian names of places, 10 trails, 13 Individual frequency, 176 Industries of pine-barrens, 17 Influence of geography on peoples and in- dustries, 12; of water, 307; of wind, 307 Inkberry, microscopic leaf structure, 274; root system of, 234 Insect-catching plants in cedar swamps, 189 Insect galls, 295 Insectivorous leaf, 263 Inter-glacial time, 6 Introduced trees, 106 weeds, 105 Investigation of pine-barren soils, 31 Ipomoea with twining stems, 311 Iris leaf form, 254 Iron-stained savanna pools, 149 Isophotic leaves, 259 Italians in South Jersey, 315 Jaccard, Paul, cited, 175 Jepson, W. L., cited, 53 Juncus militaris, submerged form, 307 pelocarpus, 5 Juniperus, juvenile form of, 312 virginiana, 90, 91, 94, 95, 96; wind-swept, 87 Juvenile forms of Juniperus, figure of, 313; of New Zealand plants, 3 12 reverted leaves of pine, 313 Kalmia angustifolia, 68, 112, 149; figure of microscopic leaf structure, 279; microscopic leaf structure, 279; root system of, 237, 238 latifolia, 70, 77, 113, 134; figure of, 239; figure of microscopic leaf structure, 279, 280; figures of sun and shade leaves, 306; mi- croscopic leaf structure, 279; mutations of, 304; root system of, 237 Kearney, T. H., cited, 267 Kiefern-Heide, 85, 86, 168 Klebahn, H., cited, 167 Kraus, G., cited, 248 Krout, Dr. A. F. K., cited, i 15 Kuster, Dr. Ernst, cited, 295 Lafayette epoch, i Lagoa Santa, 258 Lalang vegetation, 145 Lanceolate leaves, 254 Latex tubes, 262 Laurel, 77, 133, 134; microscopic leaf struc- ture, 279; root system of, 237 Law of cone and seed production in pitch- pine, 292 Layers in pine forest, 61, 62, 63 of soil, 245, 246 VEGETATION OF THE NEW JERSEY PINE-BARRENS 323 Leaf colors in autumn, 195, 209 crops, 173 forms, 252; in Denmark, 258; in Lagoa Santa, South America, 258 layer, 64 leathery, 262 mould layer, 245, 246 structures, graph of, 285; microscopic, 259; synopsis of, 283; table of, 284 Leather-leaf, 149; root system of, 239 Leathery leaves, 255, 262 Leaves as drugs, 28 deciduous, 262; glabrous, 261 ; hairy on both sides, 261; hairy on under side, 261 Lee, Francis Bazley, quoted, 14 Leiophyllum buxifolium, figure of, 236; microscopic leaf structure, 280; root sys- tem of, 236 Lespedeza frutescens, figure of microscopic leaf structure, 273; microscopic leaf structure, 272 Leucothoe racemosa in savanna, 149 Level pine-barrens, 61 Lianes, 311 Liatris graminifolia, figure of, 244; micro- scopic leaf structure, 283; root system of, 244 . ^ . , Life-giving air of pines, 310 Light and regeneration of pitch-pine, 291 Light forms of Sarracenia purpurea, 305 Limodorum tuberosum, roots and stems of, 227 Linear leaves, 253; not grass-like, 254 Linnaean species, 298 Liquidambar styraciflua, 75 List of 548 pine-barren plants, 196-208 Llanos of Venezuela, 145 Loblolly pine-barrens, 9 Local plant names, 16 Log school house at Speedwell, 19 Long, Bayard, cited, 193, 210, 313 Long Island, 5; pine-barrens, 107, 109; plants, 5 Long-leaf pine-barrens, 9 Lophiola aurea in savanna, 149 Low or wet pine-barrens, 75 Lower Plains, 151; photograph of, 152, 160, 161 Lubbock, Sir John, quoted, 252 Ludvigia sphaerocarpa with aerenchyma, Lumbering, 17 Lupinus perennis, 73, 82; color of flowers, 303; figure of, 233; root system of, 233 Lycopodium leaf forms, 253 MacDougal, D. T., cited, 313 Magnolia glauca, 134, 135; illustration of, 120; figure of microscopic leaf structure, 272; microscopic leaf structure, 271 Maltby, Robert D., cited, 172 Manasquan River, 6 Map showing geologic history, 3 Marsh formation, 48 Martha's Vineyard, 5 Massart, Jean, cited, 259 Maturity of pitch-pine cones, 209 Maxwell, Hu., cited, 19 Mayr, Gustav L., cited, 293 McCourt, W. E., cited, 1 17 Meadow-beauty, colors of flowers, 303 Meadow Brook, L. L, valley of, 171 Measurement of rhizome of Pteris, 22 1 , 222 of species of the Coremal, i 39 Mechanic analysis of soils, 33 Megaphanerophytes, 185 Melampyrum americanum ( = lineare), root system of, 242 lineare, figure of, 242; figure of microscopic leaf struc- ture, 282; microscopic leaf structure, 282; root sys- tem of, 242 Mesophanerophytes, 183 Method of pine-barren investigation, 37 of squares, 176 Microphanerophytes, 185 Microscopic leaf structure, 239; structure of leaves, detailed, 263 Microsublimation method, 276 Milk-pea, root system of, 233 Minerals in soils, 34 Miocene, i, 2 Mixed forest at Browns Mills Junction, 99; at Somers Point, 93 pine-oak formation, 48; succession, 103 Moccasin-flower, roots and stems of, 226, 227 Moisture equivalent, 47 Moisture holding capacities of soils, 43 Molisch, Hans, cited, 276 Monotropa leaf form, 237 Months of growth, 194, 193 of preparation, 193 Mucilage, 262 Mullica River, 7 Multicipital root, 218 Mutation theory, 299 Mutations, 303 Myrica carolinensis, 88, 94, 95, 96, 149; figure of microscopic leaf structure, 268; microscopic leaf structure, 268; root sys- tem of, 228 Myriophyllum humile, 308; var. natans, 308; var. capillacea, 308 tenellum, 3 Nabalus leaf form, 253 Names, local plant, 16; of early English settlers, 14; of places, 10 324 VEGETATION OF THE NEW JERSEY PINE-BARRENS Nanophanerophytes, 185 Nantucket, 5;Coremalon, 165; heathland, 168; heathland, photograph of, 170; oceanic climate of, 166 Narthecium americanum, roots and stems of, 225 Needle leaves, 253 Neocene, i Neophytes, 107 Newfoundland, 5; flora, 5 New Jersey dune plants, 285; plant forma- tions, 85; salt marsh plants, 285 New Zealand plants with persistent juvenile forms, 312; vegetation, 308 Nichols, George E., cited, 108 Nitrogen in soils, 35 Notes on distribution of transition plants, 100 Nova Scotia plants, 5 Number of pine-barren plants, excluding pines, 180; of pine-barren plants includ- ing pines, 180 Numbers of flowers in different months, 212, 213 Numerical sequence of flower development, 212 Nymphaea (Nuphar) advena, root system of, 23 1 microphylla, 303 rubrodisca as a species, 303 variegata, 231, 303 Nyssa sylvatica, 75, 112; wind-swept, 87 Oak-bottom formation, 104 coppice formation, 48; succession, 102 heath, 168 Oceanic climate of Nantucket, 166, 169,170; of New Jersey, 170 Oekiophytes, 107 Oleoresins, 28 Oliver, F. W., cited, 176 Opposite leaves, 257, 258 Orchids in cedar swamps, 190 Origin of designation " pine-barrens," 8 Original size of pitch-pine, 17 Orontium aquaticum, 147; figure of mi- croscopic leaf structure, 265; figure of microscopic section of petiole, 266; mi- croscopic leaf structure, 265 Ortstein, 166, 167 Ortstein-Kiefer, 167 Ortsteintopfe, 167 Osmunda cinnamomea, 80, 135 Ostenfeld, C. H., cited, 216, 217 Oxypolis rigidior, figure of, 299 rigidior longifolia, an elementary species, 299 Palisade leaf, 259 of two or more rows, 260; of single row of cells, 260 Palmately compound leaves, 256 Panicum Addisonii as a variety, 301 Commonsianum as a species, 301 depauperatum, roots and stems of, 224, 225 implicatum, 5 virgatum, 90, 94, 95, 96 Parasites in cedar swamps, 190 Parasitic plants, 187; in marshes, 191 Parmelee, C. W., cited, 1 17 Patanas of Ceylon, 145 Paulsen, Ove, cited, 186 Peas, culture of, in plains soil, 162, 163; in top-soil, 169 Peat, 26; formation of, i 17 Peltandra virginica, figures of leaves, 301 Peneplain, Pre-Pensauken, 1 Pennypacker, J. L., quoted, 28 Pensauken Sound, 170; submergence, 3 Percentages of leaf forms, 258 Percolation of water, curves showing, 39; rate of, 38 Perennial herbs in cedar swamps, 190; in marshes, 191; of pine-barrens, 187; with sedge- or grass-like leaves in marshes, 191 Periods of flowering anH fruiting, 196 et seq. Periwig shoots of Pyxidanthera, 241 Periwigs, 218 Persistent juvenile forms, 3 12 Pervious soils, 250 Petersen, H. E., work of, 216 Phanerophytes, 185 Phosphorus in soils, 35 Physiognomy of heathland, 167; of savan- nas, 146 Physiography, i Phytogeographic formations, 48 Survey of North America quoted, 2 Phytophenology of pine-barren vegetation, •93 Pickerel-weed leaf forms, 302 Picking huckleberries, 27 Pinchot, Gifford, cited, 126, 153, 160 Pine-barren formation, 48, 49 vegetation as a closed forma- tion, 314; old, 314 Pine-barrens, area of, 178; at northern limit, 96; kinds of, 314; types of, 9 Pine forest at Mays Landing, 82; at edge of salt marsh, 94; in Lebanon Reserve, 81; interior at Somers Point, 93; on old fields, 89 Pine grove in Shark River Bay, 103 Pine knots, 20 Pines, xerophytism of, 249 Pinnate compound leaves, 25 Pinus caribaea, 9 echinata, 9, 50, 67 palustris, 9 VEGETATION OF THE NEW JERSEY PINE-BARRENS 325 Pinus rigida, 9, 50, 69, 75, 91, 95, 96, 112, '33. 135. '37; and its fluctuations, 300; figures of branches, 300; fig- ure of microscopic leaf section, 265 ; low trees in savanna, 152; micro- scopic leaf structure, 264; roots of, 223, 224 taeda, 9 Pitch, 23 Pitchered leaves, 257 Pitcher-plant, photograph of, 122; in sa- vanna, 148 Pitch-pine, 50, 75, 91, 135; and its fluctua- tions, 300; bark of, 98; cone production of, 286; original size of, 17; roots of, 223, 224; seed, figure of, 313; seed, produc- tion of, 286; trunk of, 98 Place names, 9; of exotic origin, 1 1 Plains, 151; age of trees of, 153; formation, 48, 151; trees of, i 52 Plant names, local, 16 Plants, additional, of white-cedar swamp, 126; as weeds, 105; found in quadrats, 180, 181; systematic list of 548 pine- barren, 196-208 Plasticity of species, 305 Pleistocene, 1 * Plow-sole, 166 Pogonia ophioglossoides, mutations of, 304; roots and stems of, 226 Poison-oak, 137 Pollen-tube formation in pitch-pine, 209 Pollinati6n of pine, 209, 291 Polygala cruciata, mutations, 304 Polygonella articulata, 308 Pond formation, 48, 143 plants, additional, 144 Ponds, 143; artificial, 144 Pontederia cordata, figure of, 302; figures of leaf variations, 302 Post-glacial time, 6 Post-Miocene, i, 6 Post-oak, microscopic leaf structure, 269 Post-Pensauken uplift, 2, 5 Potamogeton confervoides, figure of, 307 epihydrus, 307 figures of, 307 Oakesianus, 5; figure of, 307 Potash in soils, 35 Precipitation, 248 Preparation months, 195 Pre-Pensauken peneplain, i, 3 Procession of flowers, 193; of flower devel- opment, 213, 2 14, 2 1 5 Propagation of shoots, 2 16 Prostrate shoots, 310 Protozoans of soils, 161 Proximity of centers of population, 3 1 5 Pteris aquilina, 69, 112, 133; microscopic drawing of leaf, 264; microscopic leaf structure, 263; root and stems of, 200 Pyrus arbutifolia var. atropurpurea, 5 Pyxidanthera barbulata, 74, 76, 152, 310; figure of, 241 ; in flower, 75; in fruit, 75; figures of sun and shade plants, 306; flowering and fruiting of, 210; figure of microscopic leaf structure, 282; micro- scopic leaf structure, 282; root system of, 240; sun and shade plants, 305 Pyxie, root system of, 240 Quadrat method, 175 1, north side of Como Lake, 177 2, one mile east of Lakehurst, 178 3, one-half mile east of Sumner, 179 Quadrats, 60; comparison of, 180; method of, 176 Quercus alba, 91, 95, 96, 133, 135; leaf forms, 303 falcata, figures of leaf forms, 303 heterophylla, a hybrid, 313 ilicifolia, 65, 68, 69, 109, 112; fig- ure of, 229; leaf forms, 303; mi- croscopic leaf studies, 268; root system of, 230 marylandica, 63, 67, 69, 84, 89, loi, 112; figures of, 228; figures of leaves, 304; figure of micro- scopic leaf structure, 269; leaf forms, 303; microscopic leaf structure, 269; root system of, 230 phellos X Q. ilicifolia, 313 phellos X Q. rubra, 313 prinoides, 66, 67, 69, 109; figure of microscopic leaf structure, 262; microscopic leaf structure, 269; root system of, 230 prinus, 135 stellata, 63, 67, 69, 90, 96; figures of leaves, 304; figure of micro- scopic leaf structure, 270; leaf forms, 303; microscopic leaf structure, 269; photograph of, 156 velutina, 64, 67, 91, 102; leaf forms, 303; microscopic leaf structure, 270 Radial shoots, 219 Radical leaves, 254 Rainfall, 248; and pine-barren plants, 249 Rate of growth of white-cedars, 1 19 of percolation of water, 39 Rattlesnake poison, cure for, 28 Rattlesnake-weed, root system of, 242 Raunkiaer, Ch., cited, 185 Red cedar, 90, 91 gravel layer, 245, 246 maple, 133, 134 Redfield, John H., cited, 6 326 VEGETATION OF THE NEW JERSEY PINE-BARRENS Reed marsh formation, 138 Reed, William G., cited, 194 Reforestation of Denmark, 161 Regeneration of fire-denuded pitch-pines, 5>. 52 Reindeer lichen, i 52 Relative importance of plant families, 183, 184 Renvall, August, cited, 286 Resin canals, 262 hairs, 262 Response to factors, 305: to light, 305; to soil, 305 Retention of water, 43 Rhexia aristosa with aerenchyma, 311 virginica, flower colors of, 303; with aerenchyma, 311 Rhizomes, 219; as drugs, 28 Rhomboidal crystals, 262 Rhus copallina, figure of, 235; root system of, 235 radicans as liane, 3 1 1 vernix, 137 Rhynchospora fusca, 5 Rikli, Dr. M., cited, 106 Rings in pitch-pine trees, 288, 290, 291 River bank formation, 48 plants, 144 Rivers of pine-barren region, 6, 7 Roads through pine forest, 14, 74; through pine forest in Lebanon Reserve, 81 Rocky Hill, 3 Roll leaf, 263 Root classification by W. A. Cannon, 247 competition, 25 i crops, 173 distribution, remarks on, 245 systems, classification of plants by, 246 systems of desert plants, 220 Roots and stems, detailed study of, 219 as drugs, 27; and rainfall, 249 Rootstock, 219 Rosin, 23, 24 Rubble plants, 219 Run-off in pine forest, 66 Russian Jews in South Jersey, 315 Sable Island plants, 5 Sachs, Julius, cited, 293 Sagittaria graminea, 307 longirostra, figures of leaves, 301 ; figure of microscopic leaf struc- ture, 266; microscopic leaf structure, 266; variation of, 301 Sagittate leaves, 255 Salisbury, R. D., cited, 1 Salt marsh at Somers Point, 96 marsh to pine forest near Ocean Gate, 95; at Somers Point, 94 Salubrious climate of pine-barren region, 315 Sand dunes of Nantucket invaded by pitch- pines, photograph, 171 Sand layer, 245, 246 Sand-myrtle, root system of, 236 Sarothra gentianoides, 308 Sarracenia purpurea, 63; in savanna, 148; photograph of, 122 Sassafras variifolium, 69, 73, 135; figure of, 230; figure of microscopic leaf structure, 272; microscopic leaf structure, 272; root system of, 230 Saunders, C. F., cited, 143 Savanna along Wading River, photographs of, 145, 146 diagram of, 147; formation, 48, 145; physiognomy of, 146; ter- races, 147 Saw-brier, roots and stems of, 226 Saw-mill on Darby Creek, 18, 19 Saw-mills, early, 18 Schematic section of cedar swamp to pine- barrens, 1 10 Schizaea pusilla, 5; illustration of, 118; mi- croscopic leaf structure, 264; micro- sco'pic section of leaf, 264 Scirpus subterminalis, 5; floating sub- merged, 307 Sclerophyll, 263 Scrape of turpentine, 22 Scrub-oak formation, 48 Sea dunes to pine thicket, 90 islands of coast, 93 Season of growth, 194 Seasonal aspects of white-cedar swamps, 127 study of plants, 193 Sedentary species, 217 Sedges in cedar swamps, 189; of marsh formations, 191 Sedgy slough in plains, 161 Seed bed, suitable, 291 development in pitch-pine, 209 production, law of, 292; of pitch-pine, 286 Sericocarpus asteroides, figure of, 243; root system of, 242 conyzoides, root system of, 242 Serpentine barrens, 9 Settlements in West Jersey, 13 Settlers, names of, 14 Shantz, H. L., cited, 45, 47 Shear, C. L., cited, 26 Sheep-laurel, microscopic leaf structure, 279; root system of, 237 Sherff, Earl E., cited, 217 Shoemaker's wax, 24 Shoots, classification of, by Hess, 217, 218, 219 propagation of, 216; structure of, 216 VEGETATION OF THE NEW JERSEY PINE-BARRENS 327 Shreve, Forrest, cited, 2 Shrubs in cedar swamps, 189 in marshes, 191; of pine-barren formation, 187; with baccate fruits in marshes, 191; with cap- sular fruits in marshes, 191; with drupaceous fruits in marshes, 191 Silene acauHs, 3 10 Sinnott, Edmund W., cited, 264 Size of plants of Coremai, i 59 Skunk cabbage, 135 Slash pine-barrens, 9 Slough in plains, 161 Sluggish run-off in pine-forest, 66 Small fruits, 173 Smilax as liane, 31 1 glauca, figure of microscopic leaf structure, 266; microscopic struc- ture of leaf, 266; roots and stems of, 226 laurifolia, figure of microscopic leaf structure, 267; microscopic leaf structure, 267 rotundifolia, 135 tamnifolia, figure of, 303 Snow scene in pine forest, 81 Soil fertility, theories concerning, 32 layers, 245, 246; numerals, 34; proto- zoans, 161; stations, 37, 38 Soils, chemic analysis of, 34; experiments with, 35; mechanic analysis of, 33; of Coremai treated experimentally, 162; per- vious character of, 250; research on, 31 Solereder, Hans, cited, 272 Solidago stricta, figure of microscopic leaf structure, 283; microscopic leaf structure, 283 uniligulata, 5 Sorting house for cranberries, 26 Sour gum, 73 Sparganium americanum, roots and stems of, 224 Spatterdock, root system of, 231 Spatulate leaves, 254 Species, elementary, 299; Linnaean, 298; plasticity of, 305; taxonomic, 298, 299; ranging from Florida to Massachusetts, 5 Spectrum, biologic, 186 Spergularia rubra, 5 Sphaerocrystals, 262 Sphagnum, 115; uses of, 26 Spongophyll, 259, 263 Spot-bound species, 217 Spotted wintergreen, root system of, 236 Spreading shoots, 218 Spring aspect, 79 Squares, method of, 176 Staminate flower development in pitch- pine, 209 Stand of pitch-pine in Lebanon Reserve, 62 Starr, Anna M., cited, 259 Stations for soils, 37, 38 Staurophyll, 259, 263 Stem analyses of white-cedar, 1 19 Stereoscopic photograph of pine forest, 61 Still for turpentine, 24 Stone, Witmer, cited, 5, 15, 143, 156, 193, 313 Storage house for cranberries, 26 Stories of low pine-barrens, 75, 76, 77 Stream bank formation, 139 Streams of plains, 1 5 1 Strips for fire protection, 100 Structure of leaves, classification, 260; of shoots, 216 Stunted trees of plains, 152 Submerged dissected leaves, 236 Subsoil, impervious, of plains, 169 Subulate leaves, 253 Succession, diagram showing, 49; of pine- barren formations, 49 Successional formations, 100 Sucker, 219 Summer aspect of pine-barrens, 80 Sundew, microscopic leaf structure of, 272 Sunflower culture in soils from plains, 164 Swamp plants, additional, 138 Sweet-bay, 134 Sweet-fern, 83; microscopic leaf structure, 267 ; root system of, 228 Sweet-gum, 75 Sweet-magnolia, illustration of, 120 Sweet-pepper bush, root system of, 235 Symplocarpus foetidus, 135 Synecology of cedar swamps, 120; of Core- mai, 154 Synopsis of leaf structure, 283 Systematic list of 348 pine-barren plants, 196-208 Table of leaf structure, 284; showing plants in flower at different dates, 212 Talbot formation of Maryland, 3 Tansley, A. G., cited, 176, 216 Tar, 23, 24 Taxonomic species, 298, 299 Taylor, Norman, cited, 2, 4, 118, 194 Teaberry, microscopic leaf structure, 278 Tension line between pine forest and cedar- swamp, 109 Tephrosia virginiana, 72; figure of, 232; figure of microscopic leaf structure, 273; microscopic leaf structure, 273; roots, 232 Terraces of savanna, 147 Theories concerning Coremai, 1 39 Theory of mutation, 299 Therophytes, 185 Thomas, Gabriel, cited, 22 Tiresome-weed, 17 Tofieldia racemosa, roots and stems of, 223 328 VEGETATION OF THE NEW JERSEY PINE-BARRENS Tolley, Howard R., cited, 194 Toms River, 7; photograph of, 140 Torches of pine splints, 20 Toxic secretions by roots of plants of Core- mal, 160 Trailing arbutus, microscopic leaf structure, 277; root system of, 237 Trails of Indians, 13 Transitional pine-barrens, 87 Treat, Mrs. Mary, cited, 29, 78, 81, 105, 129 Tree sprouts, age of, 1 54 Trees, architectural form, 53, 54, 55, 56; cultivated for fruit, 172; cultivated in the pine-barrens, 172; in cedar swamps, 189; in marshes, 192; introduced, 106; of pine-barren formation, 186; of plains, 152; with aggregate fruits in marshes, 192; with capsular fruits in marshes, 192; with drupaceous fruits in marshes, 192; with pomaceous fruits in marshes, 192; with winged fruits in marshes, 192 Trifoliate compound leaves, 256 Trunk of pitch-pine, 98 Tubeuf, Prof. C. von, 63 Tundra, 9 Tunmann, O., cited, 276 Turf -forming shoots, 219 Turkey's-beard, 77; microscopic leaf struc- ture, 267; roots and stems of, 225 Turk's-cap lily, number of flowers, 303 Turpentine industry, 21 still, 24 Types, biologic, 185, 186 Typha leaf form, 254 Undershrubs of pine-barrens, 187; in marshes, 191 Upper plains, 151; photograph of, 156 Uses of pitch-pine, 19; of sphagnum, 26; of white-cedar, 20 Utricularia clandestina, 5; figure of, 309 cleistogama as cleistogene, 311; figure of, 309 cornuta, figure of, 309 fibrosa, figure of, 309 figures of, 309 inflata, figure of, 309 intermedia, figure of, 309 juncea, figure of, 309 purpurea, figure of, 309 subulata, figure of, 309 Vaccinium atrococcum, 137; as a species, 299; figure of microscopic leaf structure, 281; micro- scopic leaf structure, 280 caesariense as a species, 300 corymbosum, 133, 134; as a species, 299; figure of micro- scopic leaf structure, 281 ; mi- croscopic leaf structure, 281 Vaccinium macrocarpon, 5; figure of mi- croscopic leaf structure, 281; microscopic leaf structure, 281 pennsylvanicum, 69, 85, loi, 112; figure of, 240; figure of microscopic leaf structure, 281; microscopic leaf struc- ture, 28 1 ; root system of, 238 vacillans, figure of, 241; figure of microscopic leaf structure, 282; microscopic leaf struc- ture, 282; root system of, 239 virgatum, as a species, 300 Vahl, Martin, cited, 185 Valley of Meadow Brook, 171 Variable leaf forms, 256 Variation, 300 Vegetation of Hempstead Plains, Long Island, 171; of pine-barrens, general, 175; of New Zealand, 308 Vegetative propagation of shoots, 2 16 Vermeule, C. Clarkson, cited, 8, 12, 178 Verticillate leaves, 258 Vesque, J., cited, 305 Vines, cultivated, 173 Viola emarginata as a cleistogene, 31 1 lanceolata as a cleistogene, 31 1 lineariloba, 304 pedata, 304 primulifolia, 31 1 sagittata as a cleistogene, 311 Violet leaf form, 255 Vitis aestivalis as a liane, 3 1 1 Viviparous acorns, figure of, 293 ; of Quercus marylandica, 292 Vries, H. de (see De Vries) Wading River, 7; savannas, 146 Waldheiden, 86 Wallace, Dr. Daniel L., soil analyses by, 34 Wandering shoots, 219 species with epiterranean run- ners, 217; with subterranean shoots, 217 Warming, Eugen, cited, 216, 239, 252, 258, 305 Water, influence on plants, 307; retention, 43 Watering Place Pond, 143; photographs of, 143, 144 Water-lily, 144; flowering and fruiting of, 210; root system of, 232 Waxberry, 88, 149 Wax-ends, 24 Weather as a business risk in farming, 194 Weeds of pine-barrens, 105 Weevers, Th., cited, 276 West Plains, 1 5 1 Western pine-barren limit, 97 VEGETATION OF THE NEW JERSEY PINE-BARRENS 329 Wheat cultures in soils of plains, photo- graph of, 164 Whippoorwill-shoe, 17 Whitbeck, R. H., cited, 17 White cedar, 134; rate of growth of, 1 19; stem analyses of, 1 19 White-cedar swamp, colors of flowers in, 190; formation, 189; photograph of, 127 White cedar, uses of, 20 White fringed orchid, roots and stems of, 227, 228 White oak, 95, 133 White-topped aster, root system of, 242 Wicomico of Maryland, 2 Wild animals of pine-barrens, 13 character of pine-barren country, 3 14 indigo, root system of, 233 Wilting coefficient of soils, 45 Wind influence, 307 Wind-swept trees, 87 Winter aspect, 78; of cedar swamps, 129 Wintergreen, root system of, 236 Wiry leaves, 262 Wiry-stemmed plants, 308 Witches'-broom on pitch-pine, 60 Woodhead, T. W., cited, 259 Woody plants in marshes, 191 Xerophyllum asphodeloides, 77; figure of microscopic leaf structure, 267; micro- scopic structure of leaf, 267; roots and stems of, 226 Xerophytic pines, 249 Yapp, R. H., cited, 216, 217, 251 Years of cone production in pitch-pines, 287, 288, 289, 290, 291 Yellow pine, 50, 67 pine-barrens, 9 Yucca leaf form, 254 EGETATON OF THE NEW JERSEY PINE-BARRENS