Table Of ContentA. 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.
