A. Dwarf, multistemmed pitch pine in the Pine Plains, with cones open following a fire. Photo, R. Good. B. Red fox pup, in the distinctive vertebrate fauna of a nutrient-scarce environment. Photo, L. Wolgast. C. Sandy road through large pitch pines and a sedge opening, probably reflecting earlier human activity. Photo, W. Ferren, Jr. D. Tea-colored acid stream with a shallow sandy bottom, bordered by Atlantic white cedar swamp. Photo, H. Olsson. E. Crown fire in the pines sweeps across a road, a common spring event. Photo, S. Little. F. Sand myrtle, a characteristic evergreen shrub of open sandy areas, is native only in the New Jersey Pine Barrens. Photo, W. Ferren, Jr. G. Snow on the pines and an infrequent ice cover on the aquifer-fed stream. Photo, H. Olsson. H. Lake wood soil profile with a gray A horizon, leached white A horizon, and irregular i 2 organo-iron staining at about 0.5 m (1.5 ft) in the reddish B horizon. Photo, M. Markley. I. Post oak locust, a grasshopper confined to the Pine Barrens along the Atlantic coast, whose irregular population explosions may strip a forest bare. Photo, J. Lomax. Pine Barrens: Ecosystem and Landscape Edited by Richard T. T. Forman Department of Botany Rutgers University New Brunswick, New Jersey ACADEMIC PRESS New York San Francisco London 1979 A Subsidiary of Harcourt Brace Jovanovich, Publishers COPYRIGHT © 1979, BY ACADEMIC PRESS, INC. ALL RIGHTS RESERVED. NO PART OF THIS PUBLICATION MAY BE REPRODUCED OR TRANSMITTED IN ANY FORM OR BY ANY MEANS, ELECTRONIC OR MECHANICAL, INCLUDING PHOTOCOPY, RECORDING, OR ANY INFORMATION STORAGE AND RETRIEVAL SYSTEM, WITHOUT PERMISSION IN WRITING FROM THE PUBLISHER. ACADEMIC PRESS, INC. 111 Fifth Avenue, New York, New York 10003 United Kingdom Edition published by ACADEMIC PRESS, INC. (LONDON) LTD. 24/28 Oval Road. London NW1 7DX Library of Congress Cataloging in Publication Data Main entry under title: Pine barrens : ecosystem and landscape. Includes bibliographies. 1. Ecology—New Jersey—Pine Barrens. 2. Pine Barrens. I. Forman, Richard T. T. QH105.N5P56 574.5'264 79-9849 ISBN 0-12-263450-0 PRINTED IN THE UNITED STATES OF AMERICA 79 80 81 82 9 8 7 6 5 4 3 2 1 Editorial Steering Committee Throughout the preparation of this volume an Editorial Steering Committee long experienced in pine barrens research assisted the Editor. Members (listed below) helped select topics and authors, review manuscripts, provide information, and a breadth of perspective. SILAS LITTLE Principal Silviculturalist, U.S. Forest Service, Northeastern Forest Experiment Station, Pennington, New Jersey 08534 JACK McCORMICK* President, Jack McCormick Associates, WAPORA Inc., 6900 Wisconsin Ave. NW, Washington, D.C. 20015 PAUL G. PEARSON Professor of Zoology and Acting President, Rutgers University, New Brunswick, New Jersey 08903 WILLIAM A. REINERS Professor of Biological Sciences, Dartmouth College, Hanover, New Hampshire 03755 JOHN C. F. TEDROW Professor of Soils and Crops, Rutgers University, New Brunswick, New Jersey 08903 GEORGE M. WOODWELL Director, The Ecosystems Center, Marine Biological Laboratory, Woods Hole, Massachusetts 02543 * Deceased. Sponsors New Jersey Academy of Science*: a nonprofit organization of professionals and others in- terested in science, affiliated with the American Association for the Advancement of Science, with the objective of improving the quality and awareness of science in New Jersey. OFFICERS OF THE NEW JERSEY ACADEMY OF SCIENCE President Executive Secretary Ralph E. Good M. Lelyn Branin Past President Recording Secretary James W. Green Mary A. Leek President-Elect Treasurer Albert F. Eble James A. Quinn William L. Hutcheson Memorial Forestt: a research facility under the care of Rutgers Univer- sity dedicated to the protection of a primeval woods, and the fostering of research in ecology, botany, and related disciplines. OFFICERS OF THE WILLIAM L. HUTCHESON MEMORIAL FOREST Director Advisory Committee Richard T. T. Forman James E. Gunckel Edmund W. Stiles Honorary Advisors to the Director Charles F. Leek Benjamin B. Stout Helen Foot Buell James A. Quinn Herbert T. Streu John A. Small (Deceased) Nathan M. Reiss John C. F. Tedrow *Hill Center, Rutgers University, New Brunswick, New Jersey 08903. fDepartment of Botany, Rutgers University, New Brunswick, New Jersey 08903. List of Contributors Numbers in parentheses indicate the pages on which the authors' contributions begin. LEE ANDERSON (169), Academy of Natural Sciences, Nineteenth and the Parkway, Philadel- phia, Pennsylvania 19103 JOHN W. ANDRESEN (283), Faculty of Forestry and Landscape Architecture, University of Toronto, Toronto, Ontario, Canada JAMES E. APPLEGATE (25), Department of Horticulture and Forestry, Cook College, Rut- gers University, New Brunswick, New Jersey 08903 JOHN THOMAS BALLARD* (133), Biology Department, Brookhaven National Laboratory, Upton, New York 11973 HOWARD P. BO YD (505), Oak Shade Road, Tabernacle RD7, Vincentown, New Jersey 08088 JOHN W. BRAXTON (373), Department of Botany, Rutgers University, New Brunswick, New Jersey 08903 HELEN FOOT BUELL (425), Box 234, Middlebush, New Jersey 08873 ROGER CON ANT (467), Department of Biology, University of New Mexico, Albuquerque, New Mexico 87131 JOHN F. CRYANt (265), Department of Entomology, Cornell University, Ithaca, New York 14853 DANIEL L. DINDAL (527), College of Environmental Science and Forestry, State University of New York, Syracuse, New York 13210 LOWELL A. DOUGLAS (95), Department of Soils and Crops, Rutgers University, New Brunswick, New Jersey 08903 JAMES B. DURAND (195), Department of Biology, Rutgers University, Camden, New Jersey 08102 DAVID E. FAIRBROTHERS (395), Department of Botany, Rutgers University, New Brunswick, New Jersey 08903 WAYNE R. FERREN, JR.i (373), Botany Department, Academy of Natural Sciences, Phila- delphia, Pennsylvania 19103 RICHARD T. T. FORMAN (xxxv, 407, 569), Department of Botany, Rutgers University, New Brunswick, New Jersey 08903 RALPH E. GOOD (283), Department of Biology, Rutgers University, Camden, New Jersey 08102 NORMA F. GOOD (283), Bioscience Department Service, Philadelphia, Pennsylvania 19103 ^Present address: R.R. 2, Box 9A, Wading River, New York 11792 tPresent address: The Earth and Space Museum, State University of New York, Stony Brook, New York 11794 ^Present address: Department of Biological Sciences, University of California, Santa Barbara, California 93106 xxiii XXIV List of Contributors LOUIS HAND (373), Department of Botany, Vincentown, New Jersey 08088 ROBERT W. HASTINGS (489), Department of Biology, Rutgers University, Camden, New Jersey 08102 A. VAUGHN HAVENS (113), Department of Meteorology and Physical Oceanography, Cook College, Rutgers University, New Brunswick, New Jersey 08903 CALVIN J. HEUSSER (213), Department of Biology, New York University, New York, New York 10003 CHARLES F. LECK (457), Department of Zoology, Rutgers University, New Brunswick, New Jersey 08903 F. THOMAS LEDIG (347), School of Forestry and Environmental Studies, Yale University, New Haven, Connecticut 06511 SILAS LITTLE (25, 297, 347), Northeastern Forest Experiment Station, U.S.D.A. Forest Service, Pennington, New Jersey 08534 JACK McCORMICK* (xxxv, 229), Jack McCormick Associates, WAPORA, Inc., 6900 Wisconsin Avenue NW, Washington, D.C. 20015 MARCO L. MARKLEY (81), U.S. Soil Conservation Service, 1370 Hamilton Street, Somerset, New Jersey 08873 PHILIP E. MARUCCI (25, 505), Cranberry and Blueberry Research Center, Rutgers Univer- sity, Chatsworth, New Jersey 08019 BARBARA MATSON (169), Academy of Natural Sciences, Nineteenth and the Parkway, Philadelphia, Pennsylvania 19103 EDWIN T. MOUL (425), 42 F. R. Lillie Road, Woods Hole, Massachusetts 02543 HANS OLSSON (245), Department of Plant Ecology, University of Lund, Ostra Vallgatan 14, S-22361 Lund, Sweden LINDA S. OLSVIG (265), Ecology and Systematics, Cornell University, Ithaca, New York 14853 RUTH PATRICK (169), Academy of Natural Sciences, Nineteenth and the Parkway, Philadel- phia, Pennsylvania 19103 WILLIAM A. REINERS (557), Department of Biological Sciences, Dartmouth College, Hanover, New Hampshire 03755 EDWARD C. RHODEHAMEL (39, 147), U.S. Geological Survey, National Center, Reston, Virginia 22092 EDMUND W. STILES (541), Department of Zoology, Rutgers University, New Brunswick, New Jersey 08903 JOHN C. F. TEDROW (61), Department of Soils and Crops, Cook College, Rutgers Univer- sity, New Brunswick, New Jersey 08903 JOHN J. TRELA (95), Department of Soils and Crops, Rutgers University, New Brunswick, New Jersey 08903 PETER O. WACKER (3), Department of Geography, Rutgers University, New Brunswick, New Jersey 08903 ROBERT H. WHITTAKER (265, 315), Ecology and Systematics, Cornell University, Ithaca, New York 14853 LEONARD J. WOLGAST (443), Department of Horticulture and Forestry, Cook College, Rutgers University, New Brunswick, New Jersey 08903 GEORGE M. WOODWELL (333), The Ecosystems Center, Marine Biological Laboratory, Woods Hole, Massachusetts 02543 * Deceased. Preface Bog iron accumulating in tea-colored streams, a tiny Pine Barrens tree frog an- nouncing its domain, fires sweeping briskly between streams, ships laden with cedar gliding out of rivers, dwarf forests reaching to shoulder height, silent cries of wolves long gone, nutrients flowing out to oysters and fish, spruce forests thriving during colder periods, a gargantuan lake resting in the sands, and a wilderness absorbing quiet recreation in the shadow of American cities—such events have inspired this book. Some we can see and some we can imagine. This volume integrates such phenomena into ecological systems (ecosystems). It describes and explores how a landscape, indeed a mosaic of interrelated ecosystems, such as the Pine Barrens of New Jersey (United States) works. It invites the reader to explore other possible relationships and thus fill the gaps in our knowledge. In short, the book asks questions, answers ques- tions, and raises further questions. Pine barrens, the ecosystem or vegetation type, has sandy or shallow soil with frequent fire, high acidity and scarce nutrients, and abundant heaths and crooked pines; Pine Barrens, the landscape, adds streams and swamps and other types to form the intertwined mosaic. People in the landscape provide the initial focus. From Indian activities and initial European perceptions of the land, the reader explores settlement, lumbering, fuel wood and charcoal, iron and glassworks, farming and livestock, and real estate development. Today, hunting for wildlife, blueberry and cranberry agriculture, forestry and forest products continue. Geology and Soils are portrayed through time and space. Sandy deposits, one on the other, are described sequentially, followed by the geographic distribution of geologic formations and soil types with their ecologically important characteristics. Mechanisms of soil development and dynamic mineralogical patterns provide a penetrating perspec- tive of soil. Climate, Water, and Aquatic Ecosystems follow, with water movement a common theme. The meteorologic conditions (both climatic and microclimatic), the hydrology of ground and surface water, a regional hydrologic budget, heat energy and evapo- transpiration, relationships between uplands and lowlands, stream chemistry and biology, and, finally, variations in water and nutrient flows affecting the surrounding estuary are presented. XXV XXVI Preface Vegetation Patterns traces the history of vegetation starting before the Ice Age. Specialists analyze vegetation using different approaches: community types, commu- nity classification according to a European method, and gradient analysis. The dwarf Pine Plains, the dynamic processes of succession and fire, and nutrient relationships of leaky ecosystems are analyzed. Finally, vegetation relationships with similar vegeta- tion elsewhere are explored. Plants of the Pine Barrens are briefly described or listed to aid the reader when species are mentioned, and when the book is used in the field. Pitch pine, common vascular plants, rare and endangered vascular plants, common bryophytes and lichens, and algae are explored. Animals and Animal Communities of the Pine Barrens include mammals, common birds, reptiles and amphibians, fish, and common arthropods. Vertebrate communities and soil microcommunities illustrate community relationships. The special ecological, botanical, and zoological characteristics and the remarkable geologic, hydrologic, and soil phenomena of the Pine Barrens have piqued the curios- ity of many, and continue to draw visitors from all continents. By describing these characteristics in the context of ecosystem interrelations, I hope a better understanding of what is really unique and special, and why, will evolve. Several important books on Pine Barrens provide a solid background. The annual reports of the New Jersey Geological Survey from the late nineteenth and early twen- tieth centuries contain several major works of ecological significance, including those of the noted botanist Nathaniel Lord Britton cataloguing the plants of New Jersey,1-2 and America's early foresters and conservationists Gifford Pinchot, John Gifford, and others describing the forests of New Jersey.3,4 From 1908 through 1911 the naturalist Witmer Stone published treatises on mammals, birds, and plants5,6'7 which even today remain standard manuals. Shortly thereafter, the plant geographer and perhaps the first of the major ecologists of eastern North America, John W. Harshberger, published an ecological vegetation analysis of the Pine Barrens.8 In the 1960's H. C. Beck and John McPhee movingly portrayed the people of the Pine Barrens as they interact with one britton, N. L. (1881). "A preliminary catalogue of the flora of New Jersey." N. J. Geol. Surv. pp. 1-233. 2Britton, N. L. (1889). "Catalogue of plants found in New Jersey." N. J. Geol. Surv., Final Rept. State Geol. 2, 25-642. 3Pinchot, G. (1899). "A study of forest fires and wood production in southern New Jersey." N. J. Geol. Surv., Ann. Rept. State Geol. for 1898 (Appendix), pp. 1-102. 4Gifford, J. (1900). "State reports on the forestal conditions and silvicultural prospects of the Coastal Plain of New Jersey, with remarks in reference to other regions and kindred subjects." N. J. Geol. Surv., Ann. Rept. State Geol. for 1899, Rept. on Forests, pp. 233-318. 5Stone, W. (1908). "The mammals of New Jersey." N. J. State Mus. Ann. Rept. for 1907. pp.33-110. 6Stone, W. (1909). "The birds of New Jersey." N. J. State Mus. Ann. Rept. for 1908. pp. 11-347. 7Stone, W. (1911). "The plants of southern New Jersey, with especial reference to the flora of the Pine Barrens and the geographic distribution of the species." N. J. State Mus. Ann. Rept. for 1910. pp. 23-828. 8Harshberger, J. W. (1916). "The Vegetation of the New Jersey Pine-barrens. An Ecological Investiga- tion." Christopher Sower Co., Philadelphia, Pennsylvania. 329 pp. Preface xxvii another and with their environment.9,10'11 Jack McCormick then published a useful preliminary ecological inventory of the Pine Barrens, summarizing vegetation charac- teristics and certain animals and plants of biogeographic interest.12 Numerous original articles underlie our understanding of the ecology of the Pine Barrens. Together these constitute an exceptionally diverse and scattered literature, written by a remarkable proportion of well-known scientists in ecologically related fields, whose interest was stimulated by the Pine Barrens. A picture of pine barrens as an ecosystem emerges which complements and enriches the pioneering work on a New York oak-pine forest by George M. Wood well and colleagues. A further objective here is the portrayal of an ecological mosaic. In this, species, energy, and nutrients are distributed in patches of varying sizes and shapes, and move among the patches. The scale is a landscape, that is, the entire Pine Barrens. Components, such as geologic formations, soil, water, vegetation, and animal com- munities, also form patches, which are sometimes spatially congruous. Interactions among patches are frequent. Furthermore, the presence and distribution of patches themselves change with forces such as soil development, succession, fire, pest out- breaks, and human activities. The dynamics of the mosaic as a whole integrates the structure, interactions, and change of individual patches. Understanding the principles of ecological mosaics will not be easy as they draw upon theory from ecosystem structure and function, island biogeography, scale, patchiness, and evolution. How- ever, the concept promises to become an important challenge in ecological theory. Too, the focus on patches of species, energy and nutrients in the landscape, and their movement between patches, has important consequences in management of the global landscape. Time as a perspective recurs through the book. We see the Pine Barrens today in terms of certain animals, plants, vegetation, and stream characteristics as described herein. But when our grandparents were about our age, John Harshberger tells us,8 "... it is rare to find trees as tall as 15 meters (50 feet). The tree as usually found ranges in height from 6 to 12 meters (20 to 40 feet)." His map also shows extensive cultivated areas and savannas in the Pine Barrens. Today those larger tree sizes might be almost doubled, cultivated areas are very limited in extent, and savannas have largely disappeared. We wonder what the area was like eons ago, or centuries ago. Too, we wonder how much it changes in our lifetime, or simply from year to year. Inevitably we wonder what it will be like in the future. Partial answers are suggested as the book describes several natural processes of change. Today in the nearly 550 thousand hectare (1.4 million acre) Pine Barrens region we see two large military bases, Fort Dix and Lakehurst Naval Station; five large state 9Beck, H. C. (1961). "Forgotten Towns of Southern New Jersey." Rutgers Univ. Press, New Brunswick, New Jersey. 278 pp. 10Beck, H. C. (1963). "More Forgotten Towns of Southern New Jersey." Rutgers Univ. Press, New Brunswick, New Jersey. 338 pp. nMcPhee, J. (1967). "The Pine Barrens." Ballantine Books, New York. 173 pp. 12McCormick, J. (1970). "The Pine Barrens: A preliminary ecological inventory." N. J. State Mus. Res. Rept. 2, Trenton, New Jersey. 103 pp.