The Pattern of Influence of Individual Forest Trees on Soil Properties Author(s): Paul J. Zinke Source: Ecology, Vol. 43, No. 1 (Jan., 1962), pp. 130-133 Published by: Ecological Society of America Stable URL: http://www.jstor.org/stable/1932049 Accessed: 27-05-2015 20:45 UTC REFERENCES Linked references are available on JSTOR for this article: http://www.jstor.org/stable/1932049?seq=1&cid=pdf-reference#references_tab_contents You may need to log in to JSTOR to access the linked references. Your use of the JSTOR archive indicates your acceptance of the Terms & Conditions of Use, available at http://www.jstor.org/page/ info/about/policies/terms.jsp JSTOR is a not-for-profit service that helps scholars, researchers, and students discover, use, and build upon a wide range of content in a trusted digital archive. We use information technology and tools to increase productivity and facilitate new forms of scholarship. For more information about JSTOR, please contact support@jstor.org. Ecological Society of America is collaborating with JSTOR to digitize, preserve and extend access to Ecology. http://www.jstor.org This content downloaded from 132.174.254.159 on Wed, 27 May 2015 20:45:06 UTC All use subject to JSTOR Terms and Conditions 130 Ecology, Vol. 43, No. 1 REPORTS cache of seeds which were later identified as C. album "just like a jar," according to the finders. The top of (Fig. 1). The seeds were in a circular, straight-sided the cache was roughly level with the lower portion of the hole about 15-17 cm wide and 20-22 cm deep that was 10- to 12-cm-deep occupation layer. The finders stated * 0 .. - that the soil layer over the cache was intact with no evidence of former burrows or other disturbance. The 4-5 liters of seeds were all as clean as that in Figure 1. Unfortunately, the finders did not recognize the possible significance of the find and it was only as an afterthought that they retained about one cupful of the material. There was no evidence of a container and no photographs were obtained in situ. The seed sample consisted of empty hulls but with vestiges of a primary root adhering to many. The seed coats, a glossy black color in fresh material, were dull and lusterless. Indications were that the seeds had germinated and the primary roots and cotyledons had later disintegrated, while the hulls were preserved intact. In view of the widespread use of C. album as a food plant by primitive peoples, the apparent man-made characteristics of the storage hole, and the location of the cache in relation to a former occupation site, it is suggested that this find provides further evidence of the gathering and use of C. album by the Blackfoot Indians in the prehistoric period. *1 UX. ...... . ..j............. REFERENCES FIG. 1. Seed of C.eonopodium album from "Ross" site: Top-showing condition of sample upon recovery, and Bottom-close-up of individual seeds. Blankinship, J. W. 1905. Native economic plants of Montana. Mont. Agr. Exp. Sta. Bull. 56. Budd, A. C. 1957. Wild plants of the Canadian prairies. Can. Dept. Agr. Publ. 983. Dempsey, Hugh A. 1960. Personal communication. Forbis, R. G. 1957. Some late sites in the Oldman River region, Alberta. Natl. Museum Can. Bull. 162. Contributions to Anthropology. Helbaek, H. 1960. Comment on Chenopodium album as a food plant in prehistory. Berichte, Des geobotanishen Institutes der Eidg. techn. Hochschule 31: 16-19. Moss, E. H. 1960. Flora of Alberta. Univ. of Toronto Press. . 1961. Personal communication. THE PATTERN OF INFLUENCE OF INDIVIDUAL FOREST TREES ON SOIL PROPERTIES1 PAUL J. ZINKE University of California, School of Forestry, Berkeley, California INTRODUCTION The soil under the influence of a forest develops properties that vary spatially with relation to the location of the trees. This variation in soil properties is frequently reflected in the distribution of the various species of the ground flora. The amelioration or degradation of the forest soil takes place with each tree as a center of influence. This paper reports the patterns of soil properties which develop under individual trees with examples drawn from work in the forested areas of California. The literature of forestry and soil science has frequent 1 Presented at the IX International Botanical Congress, Symposium on the influence of forest vegetation on Montreal. the degradation and amelioration of soils. August, 1959. reference to the effect of different species on soil properties but there are relatively few papers dealing with the spatial variation in these soil properties relative to individual forest trees. Among these papers a classic is P. E. Muller's (1887) description of the occurrence of distinctive mull humus soils under each clump of oak trees cn otherwise podzolized soils of heath lands in Denmark. Jamison (1942) noted that certain soil properties varied systematically with distance from individual orange trees in orchards in Florida. Harradine (1954) observed that in California forests where Pinus ponderosa Laws. and Quercus kelloggii Newb. were mixed there was less soil nitrogen in the open than under the influence of the pines or oaks. Fireman and Hayward (1952) found a distinctive pattern of occurrence of exchangeable sodium under various shrub species in the This content downloaded from 132.174.254.159 on Wed, 27 May 2015 20:45:06 UTC All use subject to JSTOR Terms and Conditions Winter 1962 Escalante desert in Utah. Sodium was high under Sarcobatus vermiculatus (Hook.) Torr. and low under Arternesia trideiitata Nutt. Muller and Muller (1956) noted that in desert areas the occurrence of herbaceous ground vegetation was associated with the influence of individual shrubs on the soil. The general pattern apparent from these studies is that the individual plant is a center from which soil properties may vary in a predictable manner. This paper reports an evaluation of these effects by sampling soils systematically with relation to individual forest trees in California and determining the properties of these soils. VARIATION UNDER A SINGLE Pinus cantorta The effect of a single tree on patterns of soil properties was evaluated by sampling under a 45 yr old shore pine (Pinus contorta Dougl. ex Loud) growing on a sand dune area one mile inland from the Pacific Ocean near Cresent City in northwestern California. The mineral soil was sampled to a depth of 2X2 in. at distances of 1, 4, 8, 12 and 16 ft from the tree on 4 transects at right angles to each other. The tree had a crown proj ection radius of approximately 12 ft. Two of the sampling transects outward from the tree were parallel to the prevaling winds in the area which during clear weather come from the northwest, and during storms and moist weather come from the southeast. The other transects were at right angles to these directions. Soils were sampled for volume weight and a bulk sample. The bulk samples were analyzed for pH, total nitrogen content on all transects, and exchangeable bases on one transect. The volume weight of soil on all 4 transects increased with distance from the tree. It varied from 1.17-1.32 g/cm2 near the tree to 1.65 g/cm2 in the open sand dune. The soil volume weights on the northwest transect were 1.20, 1.27, 1.44 and 1.58 at distances of 1, 4, 8, and 12 ft from the tree trunk. The pH as determined from saturated soil pastes with a glass electrode is in Figure 1. The pH was lowest (5.7) adjacent to the trunk and rose progressively with distance from the tree to 7.2 on the open sand dune. The high pH on the open dune is due to weathering shell particles in the sand. There was a change in ground flora from a single Pyrola dentata (Sm.) Piper immediately adj acent to the tree trunk to scattered Franscria Chamissonis Less. on the open sand dune. These species are associated with acid and alkaline soils, respectively. The pattern of soil pH as expressed by the isolines of pH (Fig. 1) indicates an effect of the southeast wind, the moisture bearing wind. The nitrogen content of the soils declined with distance from the tree. The highest value was 0.0286% near the tree; the lowest was 0.0025%o in the open sand. Note in Figure 2 the skewed nature of the distribution of nitrogen about the tree, oriented as if by the direction of the prevailing storm winds at the site. The exchangeable bases were determined for the transect extending south from the tree trunk. The exchangeable cations decreased with distance from the tree (Fig. 3); however the divalent exchangeable cations (Ca + + + Mg++) decreased up to the edge of the crown canopy but then rose rapidly with increasing distance outward into the adjacent barren dune, apparently because of the effect of shell material in the open sand. The exchange capacity of the soil, largely derived from organic matter in this sandy soil, declined with distance outward from the tree, continuing low outside the influence of the tree crown. The quantity of exchangeable monovalent cations (K + + Na +) deSOIL 131 REPORTS dined with distance from the tree, becoming very low beyond the influence of the tree. There is a distinct pattern of surface soil properties about this tree, and it might be postulated that in the absence of external variables such as wind or steep slope the properties of the soil under the influence of forest trees will develop in a symmetrical pattern around each tree. Thus each tree has an influence circle roughly proportional to the size of the crown area projection on the soil surface. The tree has a maximum influence under the crown canopy and the influence decreases outward from the tree. To test the generality of this conclusion transects extending radially outward from mature forest trees in various areas in California were sampled. Radial variation of soil properties about mature forest trees The general pattern of soil properties under several mature individuals of Pseudotsuga inen-iesui (Mirb.) *65N 70 6~ 0 062~ SNNW FIG. 1. The pH of surface mineral soil under of Pinus contorta. Isolines and recorded values are indicated. Wind rose (arrow flying with the indicates relative proportions of wind directions area. a tree of pH wind) in the 'IV#,~~~~~~~~' Ass~~~~~A FIG. 2. The nitrogen contents (per cent by weight) of the surface mineral soils under one Pin/us contorta tree. This content downloaded from 132.174.254.159 on Wed, 27 May 2015 20:45:06 UTC All use subject to JSTOR Terms and Conditions 132 Ecology, Vol. 43, No. 1 REPORTS Franco, Pinus ponderosa Laws., and Libocedrus decurrens Torr. were determined at locations throughout Calif. The trees were all mature (approximately 200 yrs old). Sampling of soils was carried out in surface mineral soils to a depth of 2Y2 in., usually at distances of 1, 4, 8, and 16 ft from the south side of the tree trunk. The outer sampling point was always beyond the influence of the crown canopy, its edge being between 8 and 16 ft. A series of relationships between soil properties and distance from the tree were developed (Fig. 4). The pH associated with an individual tree was almost always lowest near the tree trunk, increasing with distance outward from the tree. The nitrogen content of the soil under a tree was generally lowest near the tree trunk, increasing to a maximum at 4-6 ft out, and declining or increasing with further distance outward depending upon the nature of the adjacent vegetation. The high ~~~~~~~~~~~Capacity nitrogen associated with tree no. 1 was occasioned by sampling under an adjacent Ceanothus integerrimus H. & A. bush which is a nitrogen-fixing plant. Generally the amount of exchangeable bases was low near the tree trunk, increasing with distance to a maximum, declining Q~~~~~~~' Nat with distance beyond the influence of the crown canopy. The soil exchange capacity followed a similar trend. 3.0 2.0 .2 o E C S_ w 0.0 , I * ' Distance from tree Reet) FIG. 3. Exchangeable cation composition of surface mineral soil in the transact extending to the southeast of the single Pinus contorta. 7 Z25 /w _--;> r 5 ( E..~40., 56V 4 8 8 Reasons for the characteristic pattern of soil properties under individual trees The major reason for these patterns is the difference between the relative influence on soil properties of bark 2 ,e8 10 16 ~ 0 ~ 12 16 0 (~ ,~b go 25 4 W1 15 E 5 noa ~'On o ##V2 N 0 8 1 12 16 0 ale w~~~~~~~~~~~~~ 922 a 12 ~~~~~.150 E 0%35 4 ,'"3 j % i, ' )5 ~~~~~E p 0 4 81216 p10p 0 4 81216 Distance from Tree (feet) FIG. 4. The variation of soil properties (pH, per cent nitrogen, total exchangeable bases, and cation ex change capacity) with respect to distance from old forest trees. Numbers 1 and 2 are Pinus ponderosa; 3 is Libocedrus decurrens; and 4 is Pseudotsuga menziesii. The latter (3, 4) are from the Sierra Nevada just west of Yosemite Park, 1 is from the Cuyamaca Mts., and 2 is from the San Bernardino Mountains in Calif. This content downloaded from 132.174.254.159 on Wed, 27 May 2015 20:45:06 UTC All use subject to JSTOR Terms and Conditions Winter 1962 133 REPORTS litter, leaf litter, and no litter in the opening between trees. Compared to leaf litter, bark litter is usually very acid, low in bases and results in a mineral soil that is acid, low in bases, nitrogen and carbon. Leaf litter is generally higher in base content and nitrogen and results in a soil that is higher in pH, base content, and nitrogen content. In the adjacent opening without leaf litter the soil becomes lower in exchangeable bases and nitrogen. A nearby tree or bush may alter this effect by eliminating or affecting the open area. In addition to the variation in litter properties the quality of the rain water entering the soil may differ due to source as stemflow from the tree trunk, drip from the foliage or unimpeded throughfall and this would be reflected in soil property Thus a general, predictable soil pattern differences. The tree is about individual mature trees develops. surrounded by an inner ring of bark litter, an outer ring of leaf and twig litter, and the adj acent marginal effects of an opening or the corresponding influence ring of a nearby tree. A characteristic distribution of soil properties in the forest develops from this pattern (Fig. 5). associated with individuals of various species must be carried out with regard to these variation patterns. These patterns may be as distinctive as the depth functions or profiles of soils. Thus to demonstrate the difference in influence of various species it is wise to sample comparable distances from the tree trunk. The overall effect of a unit area of the forest on soil properties would best be determined as a summation of the individual influence patterns which occur in the area and the openings outside these areas. The denser the forest becomes the more these influence patterns merge. The older the trees the more definite the patterns will probably be. A pattern should subsist for a period following the death of a tree. The shape of these patterns will tend to be circular, but this will vary with steepness of slope, direction and intensity of prevailing winds, and shape of the tree crown. The magnitudes of the soil properties within the patterns should be functions of the soil forming factors-climate, organisms, topography, parent material and time as defined by Jenny (1941). The development of knowledge of such patterns as they occur in a forest as a function of the species present when combined with percentage cover by species in the forest should aid in determining the role of various species mixtures in soil formation, as well as in the formation of the soil-vegetation system. SUMMARY The pattern of soil properties under single forest trees is generally developed with radial symmetry to the tree, varying with distance from the tree trunk so that there is a systematic change in pH, nitrogen content, exchangeable bases, and exchange capacity and volume weights. The general pattern of this variability is due to the difference between the effect of bark litter, leaf litter, and the adjacent opening or neighboring tree. The patterns obtained are predictable in a given forest region. REFERENCES .open FIG. 5. The hypothetical pattern of influence circles on soil that would develop under an open mixed conifer forest of Pinus ponderosa (Y), Pseudotsuga menziesii Blank indicates (D), and Libocedrus decurrens (I). bark litter effect, cross hatched indicates leaf litter effect, with properties depending upon species, and stippled indicates open rainfall leaching effect. DIscuSSION The presence of distinct patterns of soil properties in the forest centered in radial symmetry about each tree means that sampling of soils to correlate soil properties Fireman, M. and H. E. Hayward. 1952. Indicator significance of some shrubs in the Escalante Desert, Utah. Bot. Gaz. 114: 2 pp. 143-155. Harradine, F. H. 1954. Factors influencing the carbon and nitrogen levels in California soils. Ph.D. Thesis, Univ. of California, Soils Dept. Jamison, V. C. 1942. The slow reversible drying of sandy surface soils beneath citrus trees in central Florida. Proc. Soil Science Soc. of America. 7: 3641. Jenny, H. 1941. Factors of soil formation. McGraw Hill, New York. xii + 281 pp. Muller, P. E. 1887. Studien uber die natfirlichen Humusformen und deren Einwirkungen auf Vegetation und Boden. vii + 324 pp. Julius Springer, Berlin. Muller, W. H. and C. H. Muller. 1956. Association patterns involving desert plants that contain toxic products. Am. J. Bot. 43: 354-355. 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