$.92»... . rAv. «V .. . A. , fikvnhyfvvrq.‘ o . V . , A A 0.1"“. . . A .. r 3..” Agavflflnf . J )4. 13 a “4...: . éol. QA AtA; . ux..r3!§p A1 . 1041 Lc‘unvvv‘vcxv. M'AVOHHr-vYNJAIUJAIVIOI as. .i _ . , a, . NW. . n.....A...u...n.... ...: a. :33. g... (:2... L . at? “$99. A .60» «AB til”. .‘Hflhat‘u fflc. . A, O ..............s ............rz¢....n..- : . .fiui .. a . . ,. .. . A. 1 njr .. Z . . . . . .. u. A ‘ .zwwfik .. . . I. V v. a .3 . .vAv-c PW .‘xuta‘cr I t w .lorv t n It ‘4 c- 1. N Q . 70. d n 00. . ..I¢I¢Av ‘01 Q. ,4. . A . 1:1 . , A. a... Huh/I. 9 INCHES (AFTER BRUNNSCHWEILER , I962) FIG. 7 ~ MEAN ANNUAL SNOWFALL (AFTER BRUNNSCHWEILER. |962I ' 15 I Writ—8*— i MEAN DEFICIT ‘ Dry 50% of Years OD . ”8 . f—* I E30-50mm. (AFTER MESSENGER, I962) 16 _-, ___._n_,,l : FIG. 9 IMEAN DEFICIT; Driesi 2 Consecuiiye Years ,5” [:3 BO-IOOmm DIOO- IZOmm I AFTER MESSENGER, I962) 1? elongation continued longer in years of ample soil noisture. According to Zahner (1963), when severe soil noisture deficits do not occur, conifers usually continue noderately rapid canbial growth throughout th entire season; in forested northern latitudes, it can be through Sept-her. He also states tint it is col-non for rapid canbial growth to resune duriu late growim season raiw periods following a W. concerning oaks, hosever, the patterns of terainal growth seen to be different in tint cessation of shoot elongation seen to be inde- pendent of scisture ceditiom, and usually occurs within two mnths after initiation (honor and Kozlowski, 1960). These authors cite an instnnce in the Missouri Ozarks where black oak, white oak and post on]: were reportedtolnveaheightgrouthseasonofonlymdays. Asturw Ids by Boggess (1956) in southern Illinois indicated that shortleai pine will resune diameter growth following a nid-groning season drought 'hlreas white oak will not. Black oak ani white oak are the min competitors with white pine jnPodnoi. Zone I. Sincethese oakspecies dcnotrespondto late growing "‘80:: soil noisture increases to the sane «tent as do pines, the meat. of the Podsol Region no; he 1... favorable fer then than for the Pines. In Newaygo county (within Podsol Zone I) the writer he observed ‘Virsinotondorrhit. pine and theabove-nentioned oahn onasand soil u'hichthe pines towerto heights 1§to 2 use.” greatas those of “*0 ssh. It so.- lihsly thot this donimnce, coupled with a long 11:.- lpan and a great old-age fire resistance, would enhnce the probability °f «hi-ring tho establishnent and survival of notnroi reproduction of “the ping. II! 18 The southern boundary of Podzol Zone II also marks the southern linit of forests dominated by red and white pines. This boundary is also coincident with the northern limit of Brunizem in Michigan. Although these linits say all be controlled by some factor(s) not yet analysed, along the western half of the boundary they are appronnated by the 1.00 - isc-defieit line of the driest two consecutive years (based on records fron 1929 through 1950; see Figure 9). Along the easternhalfofthe Zone I-Zone IIboundaryneanannualsnowfalland sntersurplusvalues arequite lowandmananmanEvaluesarenore similar to those of the Grey-Bram Podsolic Soil Region than those of the Pcdsol Region. Tint drought and high t-peratures could feasibly be limiting factors for red pins can be interpreted from the fact that miller droughts and high surface soil tenperatures frequently kill or injure young red pine seedlings (Rudolf, 1957). However, it is also POIsilble that red pine is umble to compete successfully for light with the sonswhat sore sheds-tolerant oaks which are abundant on all saw '01-'18 south of Podsol Zone II. Since Pedsol development has proceeded sore rapidly in Zone III “In inthe other sones, it is possible thatthis Ins cccurredasa 1$8111“: of the greater snowfall and autunn soil. neisttn‘e recharge. In th. autum, soil temperatures decrease sore slowly than do air temper- utum, thus November finds nest soil temperatures high enough for “Wider-able chemical and biological activity. In the case of Pcdsol 2°“. III, such activity would be particularly favored because of the reletivoly heavy sntum rainfall which would partly or conpletely re- duce nay soils, particularly those with a low available water-holding “malty (see Figure 10). These noist soils scat likely rain in an f‘ (I I“ f. l9 SOIL FIG. Io MEAN STATUS‘OF AVAILABLE MOISTURE November SOIL MOISTURE 2 8U U NILLINETERS (tear-I952) SOIL MOISTURE SURPLUS > 20 MILLIMETERS (I93I-I952) ...... PODZOL ZONE * MILLIMETERS OF SURPLUS AVAILABLE SOIL MOISTURE CAPACITY 4‘7; FEET OF SOIL 20 active state throughout the winter as a result of the insulation provided by a persistent snow cover. In tin Podsol Region of the British Isles, for instance, where snowfall is abundant, January soil tauperstures at tho ens-foot depth rams from 37.II°F to 39.203 (Jen-Hu-Chang, 1958). In the Podsol Region of Michiyn, McKenzie gt 5}. (1960) found that Kaunaska sand is noist ani unfrozen under a cover of snow in two consecutive winter seasons of study (1953-510. McKenzie's study indicates that these condition favor reduction reactions, particularly in the A1 horizon. Other studies indicate that slight decomposition of soil organic nutter tahes place even at 0°C (Kononova, 1961) and that some Podsols contain fungi in an active vegetative condition up to late fall or early winter (Dashes and Van Der Drift, 1963). Relatively favorable soil noisture couIitions in late sumer and early fall could also be especially favorable for the growth of white pine and possibly other coniferous species which are present or have beenpresent duringthedevelopnnt of the soils inPodsol Zone III. In addition, the relatively low drought intensity (as indicated by conputed soil. neisture deficits) of this zone could also be an iaportant factor by increasim the edaphic range of aesoplwtic species toward sandier soils (a Hills, 1952). CHAPTER 3. BIOIIXEICAL REIATIONSHIPS Natural vmtion _i_n Podzol Region Temperate elinte Podsols are nest frequently found under coniferous forests (Robinson, 19h9) and tho southern extent of Podzol regions often coincides with the southern extent of certain coniferous forest types (Yeatch, 1932; Tall, 1932). Possibly for these reasons, some workers have deduced that northern Podzols develop under northern conifers (Kellogg, l9hl), even though there are mnerous instances of other vege- tation types occurring on Podsols. Since a nor hams type is generally conceded to be a conponent part of mtural Podsols, it seem pertinent that the following tree species in the Podsol Region of eastern North America are consonly associated with soils having this noun-o“: Jack pine (gig banksiana), red pine (m resinosa), white pine (Ems; g_tr_o_b_u_s_), black spruce (21.43;! m). white We (:12: 522:). red spruc- (Eg .1292): balsan fir (Alias balsanea), eastern henlock (gm canadensis), red ‘1’]- flss: m» mm birch (21% 2221122): 1°11" birch (& _l_a;_t_e;), Anerican beech (m Endifolia) and sugar nple (egg saccharwn) (Ra-ell. and Heiberg, 19313 hint, 1932; Donahue, 19393 Iilde, 1958 ad personal observations). Association of the latter threespecies, however, arenore co-onlyfoundonmllhunus types thanonncrhnmstypes enceptwhsre theproportionofbeechis high (no-oil and Home. 1931). Upland species growing in tho sane areas ¥ EOnJythcsehu-islayersareinclndedwhiehneetthenorhunus Ipecificatiou outlinedbynooverasd Lent (1952). (‘9 22 but found nostly on null hunus types are: red oak (Qg_er_§_u_s_ 53253), Anerioan basswood (Lilli: anerioane), white ash (Fraxinus anericana), iromod (9.2353. M), Ana-icon Eln (PM anericana), black cherry m serotim) and butternut M cinerea). Aspem (My. tremloides 2.51. Edidentata) are present throughout the region and their abundance is usually a reflection of past fires or other denature disturbances (Spurr, 1961:), thus they are usually associated with null or duff-cull hums types since nor bums types are usually degraded by fires and disturbances which open up the forest «new (Ron-n. 1935). Plies (1931;) noted that the heaviest and nest pronounced nor hum is found under pm hemlock stands. Under this hum typo, the A2 (En) horison reaches snximn thickness and cleanness of color. In addition, the H-layers (0h horisons) are here acid under pure honlock than under W other forest type studied in the northeastern states, the pH range being fru 2.5 to halo Nearly ooIanrable humus layers, however, lave beenfcand inPodsels underspruce-fir stands inNew Hanpshire (Int, 1932) and under black spruce stands in Quebec (lafond, 1958). The southern boundsry of the Podsol Region of Michigan coincides with the southern linit of the area in which white pins (3% strobus) is an ilportant conponent of the recent mtural vegetation (Vesteh, 1932; see Figure 3) and closely parallels the southern botanieal range limit of henlock in tint state (233 Bough, 1960). Podsol Zone I essentially coincides with the vegetation ”tension none" and is bounded onthe northbythe southernrange linit of redpine (see Figure 3). Ihite pinswas the only pine of ilportanoe inthis zone and it occurred Iinly in fixtures with ed: on the well-draw sites. In Podsol Zone II, 23 Jack pine Ens; banksiam) and red pine (m resinosa) were also present (Figure 3). In this zone, henlock was more prevalent (Rough, 1960) end the pines occurred in pure stands* as well as in nixtures with hrdwoods. The pure stands of pines were present alnost exclusively on sands, the nest prevalent of those being Rubicon send (see soil survey reports of Newaygo, Montcala, am Midland counties). Northern hardwoods were generally prevalent on Iodine-textured and fine-textured soils in this acne with so. notable exceptions in the northeastern port of the Lower Peninsula where white pine dominated on some clay loan and silty clay loam (Veateh, 1953, 1959 and persoml consummation); these soils belong to the Gray-Wooded great soil group (NOR-3 Soil Survey Co-ittee, 1960). D: Podsel Zone III, northem lardwoeds were more prevalent and svenoccnrredonsonewell-drained sands, while pines or oaks andpines occupied other well-drained sands. Here, the Nine-textured soils invariably snorted northern hrdloods with scattered white pines and henlcck often msent, this relatiomhip utendim on into the Upper Peninsula. new The pattern of fire-free forest succession in the lake States is saidtobefroejackpineteredandwhitepines toshade-vtolerant species suchassugarnple,balsanfirandhlackspruceinthe caseof sands (Spur, 1961;). amoral-successions]. chnges tro- jack pine te redandwhitepines issuggestedforsands innorthernlewerlichipn whilewhite spruceandbalsanfir throuhwhiteandredpinestehenleck *‘Purestands'nfertostandscolposedefatlaastmofthestated cosponent. Fun ”I nan ’\ .e 21.; and northern hardw00ds is suggested for the finer-textured soils (Kilhurn, 1957). lithin Podaol Zone III, sands occur which have (or had) pure stands of northern hardwoods on them. To take these and other sand sites into account, perhaps Kilburn's cliseral-euccessional pattern should be revised to: from Jack pine to jack pine and oaks on some sands (role of fire my be important), to red and white pines in the case of other sands and Im white spruce and balsam fir through white and red pines to hemlock and northern hardwoods on still other sands and finer-textured soils e All these sequences were probably initiated by relatively short-lived stages characterized by calciun tolerant, shade intolerant species similar to those mentioned by Olson (1958), Cracker and Inger (1955), and Wright (196b,), some of which produce easily deconpoeed pollen and therefore may not be accurately repre- sented in pollen spectra. Regardless of difficulties of interpretation, pollen studies do indicate that a spruce-fir period dominated all forested land stn'faces in Michigan following Cary glaciation (Potzger. 1916, 191:8; Wilson and Potager, 191:3; Pamelee, 191:7; see Figure 11). Even in southwestern Michigan where oaks and northern hardwoods now donimte the upland sites, spruce and fir reigned supreme for approximately 14500 years following Cary glaciation (Zunberge and Potsger, 1956). According to these findings, pine was the njor vegetation type for the succeeding 3500 years and it was not until about 5000 B.1=. (years before present) that hardwoods and hemlock entered the scene. In Choboygan County, about 250 niles further north and well within the Podzol Region, Kuburn (1957) estimtes that spruce and fir fa'ests were donimnt I. ELEV. 660 FT. FIG. II POST— GLACIA'L ELEV. SOOFT. ELEV. 900 FT. - ' PINUS- PICEA I] .POLLEN PROFILES PICEA-ABIES L 0 ME\ QUERCUS- PINUS J QUERCUS PlNUS-QUERCUS. / ‘ \~/- ousacus- NO. HDWD. PINUS- No. HDWD. PINUS- TSUGA PINUS-BETULA '~ =LENGTH OF AREAS WITHIN HISTOGRAMS THE PERCENT OF SAMPLING DEPTH REPRESENTS 26 for the first 3000 years following Valders glaciation and that pine forests were prevalent for the succeeding 1000 to 2000 years. Although no mention is nde of succession to northern hardwoods on sand, Kilburn infers that conversion to northern hardwoods and hemlock on loany sands occurred sonswhat later tun on loans, possibly from 2000 to 3000 years ago. This chronology would of course imply that coarse-textured drift of Valders age Ins supported prodonimntly coniferous forests for at least 8000 years. Pollen data are scarce for the Upper Peninsula of Michigan, but studies in northern Wisconsin (Potsger, l9I46; Wilson, 1938; Wilson and Iebster, 19th) indicate a forest succession similar to those of the northern part of the Icwer Peninsula of Michigan with the exception tint birch is outstanding in some of the Iisoonsin pollen profiles; ncrofossil data free Minnesota indicate that paper birch was the birch species which invaded the deteriorating spruce forest (Wright, 1961:), while upper level birch pollen in north central Uiscomin probably represents ninly yellow birch (putts-r, 19%). The peat sampled. by Iilson and Iebster on an outwash plain in north central Wisconsin indicates an initial white spruce marina that was quickly replaced bya pinenximnwhichpersistedto the present tine. Iathe upper one-third of the profile a definite increase in birch (probably yellow birch) an! henlock pollen was present. Two other bogs, located in areas of finer-textured soils, showed an increase in spruce near the surface, acre or less aeooupawim the birch and hemlock increase occurring in all three profiles. 2? Fraxinus (ash) was present in southern Delta County by 5720 2:250 years B.P. as determined by a radiocarbon dating on a piece of buried w.od*. The above studies suggest that oak was more prevalent in Podzol Zone I than in Podsol Zone II. They also indicate that hemlock was relatively abundant north of the tension zone whereas hemlock pollen percentages were consistently less than 10% in Podsol Zone I bogs. Ho henleck pollen was found in the bogs of Douglas County, Wisconsin (entrees northwest corner of the state) by Iilson (1938) suggesting that henlock did not play a significant role in post-Pleistocene plant succession in the less humid portions of the labs States Podsol Region. III Douglas County, only weakly developed Podaols (no dark orterde hcrisons) are found and these only on and parent nterial (MR-3 Technical Conittee on Soil Survey, 1960). According to Wilson the forest histoly of these sands was from jack pine and spruce (concurrent with Glacial lab Duluth) through jack pine an! red pine (durixg Glacial Ialnsilgonquintins) toredpine, Jackpineandoak (beginninginlahe Nipissiu tiles). The oak conponent was likely present to the greatest extent entheweaklypodsolised site (mega sand, whichis usmlly classified as a Brown Podsolio soil). Appreciable quantities of oak were ehrecteristic of the mtural vegetation (recent) on this soil. series and not chucteristis of the Podsol sands (Rubicon and Vilas series) inthisareaalthoughnerthernredoakeanbefound ensue areas of Rubicon and Vilas soils. 'Persoml co-anication free A. E. Slaughter, Geological Survey, Division of the Departnnt of Conservation, State of Michigan, at Bsoanaba. The wood was identified by Dr. Eldon l. Bohr, Depart-ant of Forest Products, Iichigan State University. Judging from the site description given by Hr. Slaughter, the tree was evidently inundated initially hy the rise of water levels from the Isles Chippewa stage to the lake Iipissing stage in the lake lichigan Basin (see Rough, 1958). f. I. l‘ 28 Vegetation gm g Associated Changes _i£ Soil Mogholcg Disoussiu the soil. types of south Sweden, Tenn (1932) is of the opinion that there Brown Forest soils (a group of soils in Europe sons ofwhichnybeequivalent tothe BrownForest soils of the U. S. and sons of which my be equivalent to Gray-Brown or Brown Podsolic soils ef tb U. 3.) are the clintically determined soils and he describes their occurrence on may different parent materials if the natural vegetation of beech and oak forests is present. Ibere, hemver, the broadleaved forest has been replaced by conifer forest or Callum heath, as often ripper under the influence of fin, Brown Forest soils develop- ing into Pedsols are found. Tm also describes the phenonenon whereby aolearlydefinedPodsolnyacquireamllhuaus layerandaless acid reaction if beech or conifers are replaced by birch. He also points out thatifbeecherspruce colonies orareplantedunderthebirohonsuch soils then a ner him will be found again. Handley (19514) cites references to the fact that on base-rich soils, European beech gives risoteanllhu-ls layerwhereas onabase-poor soil, it gives rise to a nor. Sinilar phenol-one hove been observed by Fisher (1928) and Griffiths, lartwollandShw (1930) inlIewKngland, wherewhite pinehas developed on abandoned fields. After 80 years there is alnost no vegetation under thewhitepine,andundorthethinlayerofdryneedlesthereisathiok layer efrawhumsand 'a stronglymdsolisedhorison'. Onanadjacent plethrdIoedforosthasbeendevolopisgonasiailarwhitepinoplot which led a sinilar profile. at the ties the white pine us relieved; new thereisatruenllhunuspresonh—allaocunulatedrawhtms hasnerged withthenineral seilandlesstlnna singleyear's leaffallraains on the surface. I 29 More recently Bornebusch (33.93 Handley, 19514) has described profound changes in soil norphology brought about through the influence of Quercus Elba: (introduced fron the U. S.) planted on nor humus layers produced by m glvestris and Page: melee on sandy soils in Denmrk. In 20 years the bleached A2 had bosons obscured; the nor layer had largely disappeared and us replaced by a brownish, earthworn null humus layer. Possibly related to the above studies are the findings of Nikola (195” who noted tint soil basidic-ycetes docuposed leaf litter nore rapidly the needle litter and those of Ivarson and Sowden (1959) who found tint fill-foreleg litters dooonposed more rapidly than nor-forcing litters. Although several ecqflamtions of these differential rates of decon- position have been given, the work of Iossaint (1953) indicated that the rate of deco-poutien of litter fun 9 species was directly related to the l and water-soluble Ge content of the litter. Cheeical m 4.3.9.1.: _i_n Forested Romans To obtainefullunderstanding ofsone oftheafore-eentioned phenoeon, it is becoeing increasingly apparent that a knowledge of couplete ecosystem is necessary. For sample, recent studies on forest ecosyst-a have provided useful inflation concerning the distribution of nineral eluents one they lave been initially renoved free the soil by plant reots. Betintes of the conposition of standing crops of living trees give sue notionas towhat he beeainths soil. but is set includedwhensoil anlyses are nde. The following tables represent three floristieellydifferent standing tree erops (based onweighingand saspling tb various parts of sanple trees): 30 Pounds per Acre Site Species 517 Better T—‘T—T—‘Ca Mg emu-11 Beech 110,1.90 - 31.1 121.2 261.2 39.3 stud- Birch 38,735 - 9.6 116.2 103.6 1b.? Great Fir 1,675 - 0e3 101 2o9 Oe3 Seohy Spruce 561 - 0.1 0.3 1.1 0.1 Mountains" __ Total 15171161 - HI?! 13823 37178 5575 Natural Spruce 161;, 788 - 22.2 102 .9 251.1 35.}; Stand— Fir 93,386 - 13.3 57.6 152.0 19.0 Great Birch 31, 78h - 8.1. 36.7 81.h 11.9 3.91? now-rum” no.1 2897998 '1" 53".? I973 138576 3672 6!; year .10 Scots pine 106,000 183.0 20.3 89.0 180.0 31.8"” plantation on dune sand -Bcotland*" *snnh et .1., 1961. alright 3501111, 1958 and Wk. 1959 The greater aaount of standing stock in the spruce-fir stand compared to the beech-birch stand ny be a reflection of nors efficient site utili- sation, a reflection of a differeme in the sites or silply a reflection of an innte difference due to the growth fora olnraotsristies of the spruce as coupared with the beech donimnts. The greater biomss in the spruce forest results in nore nutrient element naterial in the above- groundportionoftlnt ecosysteatlan inthatoftho hardwoodecesysten. Results obtainedbymington (1956) indicateasiailartrendinthe cases studied. The cheaioal ole-ont concentration in the vegetetive nterial, however, is also of considerable significance to the deco-positionl processes tam place in such ecosystem. For msple, dataforthothreetreecropsaboveindioatcalowerconcen- tretionofallthe elements inthe conifer stands as coaparedto the beech-birch stand. 31 To obtain a clear picture of what takes place in such ecosystems, studies need to encompass the other portions of these ecosystem as well. For instance, the various chemical elenent pools in forested terrestrial ecosystems can be visualized using a model such as the following one which also depicts the possible directions of exchange. Chemical Element Pools in Upland Forested Terrestrial Ecosystem“ Stem and Foli- Grass Roots and Available Elements Fixed and Minersl- Sub-e olum Incorporated Zone Zone Zone Zone Elennts *sizes of rectangles do not signify relative pool capacities or qmntities. The forest floor should logically provide the most constant source of biologically controlled chemical environment for the underlying horizons of the soil profile. The forest floor not only reflects the chenical element composition of the entire supra-solemn portion of the biosphere but it has features which can alter the fern of the chemical elemnts and conpounis which enter it from the atmosphere. The mjor annual chenical element contribution to forest floors, however, is from tree foliage thus forest floor differences are largely reflections of foliar differences when similar atmospheric conditions prevail. 32 Data presented by Remesov (1958) suggest that seasonal transfers of son chenical elenents fro- available element pools in the soil to aware- solue pools can retard their removal from the ecosysten by leaching. For maple, ale-inn: losses were less when vegetation development in the spring preceded spring leaching. Losses of aluminum from pine-dominated watersheds were consistently less tinn those from oak- or aspen-dominated watersheds when leaching was subsequent to the beginning of spring vegetative growth. Whether or not this difference was a result of greater alunnua uptake by pine could not be inferred from the data given. (Sore United States data suggest that white pine my taloe up acre ale-inn: than red oak, for example.) Leaching losses of calciun and potassiua were greater a__ft_e£ the resumption of biological activity in spring. Romanov attributed this relationhip to the biological conversion of calciu- froe a water-insoluble to a water-soluble fan in the forest floor. He attributed the greater amount of calciu leacMng from oak and aspen watersheds (as compared to pine watersheds) to the higher calciul content of the oak and aspen leaf litter. leaching losses in the studies of Renesov were confined to spring. According to Volubuyev (1959) Podaol region! have spring and fall leachingseasons. Onsandsoils mpodsnm IIIinllichigan, water balance computations indicate that ecosystem losses my occur in fall due to leaching (Messenger, 1962; see Figure 10). Thus the ecosysten distribution of leaclnble elouents during the fall leaching season could also influence losses therefrom. 1h nest of the Podsol Region of Michigan, precipitation exceeds potential evapotranspiration from Septeaber through April or my. Studies 61m by Koslowsld. (1960) indicate that chenical eluents in tree foliage do not increase after Septeaher and tint calciue content is at a sexism in early fall whereas 33 the spring leaching period is characterized by increasing absolute foliar contents of nost elements. If foliar contents reflect root uptake, fall leaching would not be reduced as mch by plant uptake as spring leaching would be. Elenatal Gwition 21; 33:2 Foliage 22d E33352 Based on sole of the above relationships, a number of worloers have approached hunns type and soil development from the standpoint of differ- ences in the elemental composition of tree foliage. Even though the form of the cheaieal elements in tree foliage is important in such studies (Eandley, 1951:), lost analyses have been concerned only with the concen- tration of the chenical elements regardless of their fern. Several studies of this nature have been ads on the foliage from trees which are found in the Podsol Region of North America. Since the elemental comositicn varies with leaf age, soil conditions and clinte (Kramer and KoslowslIi, 1959) the following tables are presented so that foliar comparison can be nude: (1) between different species in the sane generalaree growing on the sun or similar soils, and (2) between individuals of the cane species growing on different soil types. Tables 1and2 sunrisothe intact foliageandfreshlyfallcnlitter data from the literature which point to species differences in foliar conposition of njor nutrients. For each site the species are arranged in order of decreasing calciua concentration. These tables are not conbined because of known variation in foliar composition resulting from (1) translccationofeleuentsback intothetwigspriortoleaf fall and from (2) leaching of soluble elements during leaf senescence and follaing leaf fall. Couparisons between the two tables show a general decrease inn, Pand [withleafago thussuggesting that one 3h or both of the above types of foliar decreases have been operative. The crown position of the foliage samples analyzed for Table 2 were not “me cuposition attributable to the portion of the crown sampled (Kramer and kilowatt, m0)e Some investigators, however, have found variations in foliar TABLE 1. Geqosition of Freshly Fallen leaves from Different Tree Species Growing on Similar Soils in the Same Vicinity 5m Species %N 50a xx 7062 %P Case lab fine Basswood 0.97 3.114 - . 0.18 sand near Star Sugar Maple 1.32 2.57 - - 0.10 Island, lflm. Red Oak 0.6!. 0.96 ~ - - - (my :3 31., lhite Pine 0.53 0.97 0.17 - 0.07 1933) Red Pine 0.67 0.96 0.21; - 0.07 Jack Pine 0.59 0.63 0.18 - 0.05 Scarbore lean Sugar laple - 1.08 0.112 0.1714 0.10 sand near Iitchfield, Red Maple - 0.93 0.39 0.148 0.12 Harri-e loan Red ml: - 0.61; 0.140 0.26 0.211 sand near Litchfield, mu Pine - 0.1.5 0.36 0.22 0.15 Gem. (Scott, 1955) Sandy lean Pedael, White—cedar 0.60 2.16 0.25 0.15 0.01; Web (CW, 3‘18“ Fir 1.25 1.12 0.12 0016 0e09 1910) Rod Spruce 0.89 0.79 0.35 0.20 0.10 Hemlock 1.05 0.68 0.27 0.11. 0.0? Ihite Pine 1.1!; 0.60 0.18 0.16 0.05 Red Pine 0.69 0.58 0.35 0.18 0.07 m and silt Basswood 1.0h 3.21. 0.39 0.39 0.11: lean Gray-Brown Black Cherry 0.55 2.58 0.1;? 0.“; 0.18 Podaolic soils, Iromood 1.01 2.52 0.35 0.35 0.09 central New Ierk Ihite Ash 0.59 2.28 0.h6 0.29 0.15 (Chandler, 191:1) American Eln 0.77 2.06 0.1.); 0.32 0.15 Aspen 0.70 1085 0e36 0023 0e08 Sugar laple 0.173 1.65 0.16 0.28 0.12 Red on: 0.67 1.119 0.55 0.31 0.11 Red mp1. 0.141 1.35 0.30 0.32 0.11 Ihite Oak 0.50 1.22 0.52 0.21. 0.12 Beech 0.59 1.09 0.65 0.26 0.10 ,‘\ f. fie 35 TLBII 2. Composition of nature Foliage” from Different Species on Similar Soils in the Same Vicinity Site Species %N %Ca 51! %Mg %P DeKalb soil series Ihite ish - 2.3 - - - (no 11-9 in root Yellow Birch - 1.6 - - _ zone). Warren 0mm. Eastern Hemlock - 0.8 - - - mo (P3100. 1933) Northern Red (3]: - 0.7 - .. - “it. P1110 "' 0.5 - "' " Glacial till soil White Spruce - 1.9 - - — (no 11" in 1‘90" '01”) Eastern Hemlock - 1.1 .. .. - in Adirondacks (Plies, 33,139,, Fir .. 0,9 .. .. .. 1933) Black Spruce - 0.9 - - - Dunldrk soil series White-cedar - 2.6 - - - (lies present in root American Ela - 1.6 - .. - acne) near Ithaca, NJ. Sugar Maple - 1,1 .. .. .. (Pliee, 1933) Ihite Oak - 0.9 - - - Red Maple - 0.8 - - - Tanrack - 0.6 - - - Pitch Pine - 0.1: - - - 3111; lo:- shcnl Basswood 2.68 2.88 2.16 - 0.26 till I911 (line Ilium! Ironrood 2.01 2.62 0.96 - 0.15 1 toot or soil mace) Black Cherry 1.67 2.13 1.63 - 0.18 in central New York Yellow Birch 2.56 1.86 1.10 - 0.18 (Bard, 191:5) Ihite Ash 2.27 1.70 1.70 - 0.18 Sugar mp1. 1.81 1.55 0.78 - 0.09 American B1- 2.86 1.5!. 1.00 - 0.11: Red Oak 1.61. 1.25 1.50 - 0.11: American BS“ 2037 0097 1.00 - 0.111 Silt M 311-0131 Basswood 2.32 2.87 2.35 - 0.27 tin soil (Id-9 at about Ironwood 1.62 2.59 1.10 - 0.15 30 inch» depth) in me. Ash 1.91 1.811 1.5h - 0.28 camel HUI Ink (Bird. Sugar Maple 1.68 1.63 1.02 - 0.16 19115) Ihite Oak 2.33 1.60 1.55 - 0.211 Red mp1- 1.1;); 1.1.2 1.16 - 0.16 mm Mb 2e21- 1e21 1.39 "' Gem Red 0.1: 1.75 1.02 1.20 - 0.18 Hemlock loll 0e70 0e” " 0.18 Silt 1“- 8110111 330m 2“} 2e96 2e30 " 0033 1:111 soil (no line in Ironwood 1.99 2.1.7 1.30 - 0.19 root zone) in central White Ash 1.90 1.68 1.37 - 0.27 now Ierk (Bard, 191:5) Big-t. Aspen 2.10 1.67 1.00 - 0.26 White on: 2.06 1.1.1 1.1.6 - 0.21: Yellow Birch 1.92 1.37 1.35 - 0.25 Red Maple 1.13 1.11; 1.02 - 0.25 Sugar llple 1.50 1.11 1e2h " 0e21 Red 0a]: 2.29 0.91 1.52 - 0.25 American 33.011 2.03 0.35 1.21 "' Gel? Hdwk 1e33 0e71- 1e20 - 0018 36 Site Species 921 20a %K %Mg 51> Site #7616, Dane County, Red Oak 0.96 1.18 0.29 0.50 0.12 Iisconsin (Gerleff Sugar Maple 0.73 1.01 0.39 0.115 0.12 g 31., 1961;) Site #18:», Dane County, Basswood 1.39 1.11 0.89 0.52 0.17 23'. 21." 1965) Site #1111, Dane County, Butternut 1.79 1.11 0.82 0.72 old; 31 g_l_., 1961.) Site #18:, Dane County, White Oak 1.03 1.13 0.70 0.32 0.12 Iiscomin (Gerleff Ironwood 1.1;? l.06 0.611 0.51; 0.16 o_t s__1_.., 19611) Big-t. Aspen 1.11 0.99 1.78 0.33 0.21 Iisconsin Gerleff White Oak 2.19 0.82 0.85 0.36 0.19 23-. 21;. ) Red on: 1.97 0.75 5 .39“ 0.1.0 0.1L: Black 00.1: 1.92 0.70 0.77 0.112 0.11; Site :62, Vilas County, lhite Spruce 0.95 0.87 0.5h 0.23 0.17 wisoonsin (Gerleff Balsam Fir 1.22 0.75 0.116 0.13 0.1.3 2 20’ M) Site #55, Vilas County, White Pine 1.h8 0.32 0.5!: 0.23 0.15 Wisconsin (Gerloff Red Pine 0.95 0.23 0.1.1 0.17 0.11 33 9,, not.) *Cenifer needles of the current year were used in the Wisconsin study; it is assuned that in the other studies the conifer needles analysed were variable in age. “This value scene to be too high. ("I ,Q 37 T1313 3. Couposition of Freehly Fallen leaves from the Same Species on Different Soils in the Same Vicinity Vicinity: near Iitchfield, Connecticut (Scott, 1955) Ridgebury Scarboro Merrimac Merrimc lean Loan Loan Sand Sandy Loan (03*); null. (110*); nor (BF‘); nor (BP*)3 nor R“ %0a 0099 0093 "' '- Maple 5x 0.117 0.39 - - $112 0.1.6 0.1.8 .. - 5? 0.11 0.12 .. - Ihite 50. 0.66 0.69 0.11. 0.149 Pine %1 0.145 0.113 0.38 0.37 $Mg , 0.33 0.32 0.23 0.31 $13 0.10 0.11 0.11; 0.11 Vicinity: New Iork (chandler, 191:3) Sandy Leas Silty Clay Loan Gray-Brown Podsol Podzolic Soil Ihite %ca 0.60 0.60 513 0.16 0.21 %P 0.05 0.07 *ss=srsysydranorphis;m=aunic01sy;Bp=Brownpodxonc 38 TABLE 1;. Composition of Mature Foliage from the Same Species on Different Soils in the Same Vicinity Vicinity: central New York (Bard, 19115) Silt Lean Glacial Silt Loan Glacial Silt Loan Glacial Till Soil With Till. Soil With Line Till Soil With Line Within 1 Foot at About 30 Inches No Line in Root of Soil Surface Below Soil Surface Zone in 2.68 2.32 2.1.1. Basswood $0.. 2e88 2e87 2e96 %K 2.16 2e35 2030 z? 0.26 0.27 0.33 %N 2.01 1.62 1.99 Ironnod 50s 2.62 2.59 2.17 “ 0e96 1.10 1e30 52 0.15 0.15 0.19 %N 2.27 1.91 1.90 Ihite 50: 1.70 1.81. 1.68 Ash %K 1.70 1.51. 1.37 %P 0.18 0.28 0.2? $11 - 2.33 2.06 Ihite %Ca - 1.60 1.111 0.1: $2 - 1.55 1.116 %P - 0.21. 0.21. am 2.56 - 1.92 1011“ $0!- 1.86 " lea? Birch 52 1.10 - 1.35 52 0.18 - 0.25 in - 1.1.1. 1.1.3 Rod 20. - 1.112 1.11. 5P - 0.23 0.25 in 1.81 1.68 1.50 Sugar 50. 1.55 1.63 1.11 mp1. :1: 0.78 1.02 1.21. 12 0.09 0.16 0.21 in 1.61. 1.75 2.29 2.11 50. 1.25 1.02 0.91 on: 51: 1.50 1.20 1.52 52 0.11. 0.18 0.25 in 2e37 2.21 2e03 American 10. 0.97 1.21 0.85 - Beech St 1.00 1.39 1.21 5? 0.11. 0.11. 0.17 in - loll 1033 Eastern m - 0.70 0e71- Huloch xx - 0.90 1.20 11’ " Gel-8 0.18 39 The studies nude on foliar N suggest that the freshly fallen foliage of Jack pine, red pine, white pine and northern red oak have similar low contents as compared to sugar naple and basswood when all six species are graing on sand. 0!: better sites, white pine seems to have considerably higher contents than red pine and similar values to those of hemlock (Table 1). Mature foliage studies in Wisconsin also indicate that white pine contains a higher percentage of foliar N than does red pine (Table 2). The ntm-e foliage of hemlock, as indicated in Table 2, contains lewsr percentages of N than any of the associated hardwoods. Since the age of the hemlock foliage is not known, interpretation of this cens- parison is difficult. Variations with age may not be great, however, since Gerleff gt _a_1. (19611) give a percentage of 1.21 for current nature foliage while the freshly fallen litter value reported by Chandler (19113) is 1.05. The values reported by Bard for uture hemlock foliage of unknown age are 1.11 and 1.33. The native foliage data in Table 2 also indicate that white ash, yellow birch and beech have higher foliar N values than sugar nple. m acid sites, northern red oak also has higher percentages at 3 than sugar nple. Basswood tends to have high values of foliar N wherever it occurs. Studies nde on foliar Ca from different species growing on the sale or sieilar sites, Table 1, suggest that freshly fallen Jack pine needles hve a lower content than either red or white pine. The foliages of pine, hemlock and beech consistently have lower Ca concentrations than associated hardwoods emcept red oak. The foliages of basswood, iromrood, butternut, black cherry, white-cedar and white ash contain relatively high concentrations of Ca. ,1 ho Table 2 indicates that the mature foliages of basswood, white ash and black cherry comistently have relatively high concentrations of potassium. Ihen associated with hardwoods, the nature foliage of hemlock has potas- sin «to. sinilar to those of its associates except that basswood foliage consistently has higher values. The lardwoods in Tables 1 and 2 have foliar magnesium values which vary along a ccntimous gradient with butternut, black cherry, ironwood and basswood at the high end and white oak, aspen and beech at the low end. A relatively low range of foliar Mg values is indicated for all of the conifers. Despite the paucity of site—mte conparisons between hard- woods and conifers, it is interesting to note that the highest value reported for the conifers is the sane as the lowest value reported for the hardwoods. Site-ate comparisons made by Ovington (1956) in Great Britain indicate that with the exception of certain oak species, hard- woods contain higher foliar Mg concentrations than do the conifers. Site-nte comparison indicate that the foliages of basswood, black cherry and white ash contain relatively high concentrations of phosphorus am that the foliages of beach and pines lave low concentrations of that slant. Several species in the foregoing studies show evidence of a foliar nutrient element response to site. Most of the hardwood species studied by Bard, Table 14, show increasing foliar N concentrations with decreasing depth to carbonates. Northern red oak and henlock, however, show decreasg contents of foliar N with decreasing depth to carbonates; this trend-.ybea general one for these species sincethe-zimrecorded value for northern red oak foliar N was obtained for leaves collected frcna northernred oakplantation onanacidsand inhgland. The average $1] there was 2.87, for higher than the values for an of the I. I. hl seven species of evergreen conifers planted on the same site but similar to the values for three other members of the Fagaceae family on the same site (1133 Ovington, 1956). Data presented by Chandler (l9h3) was thought by that author to confin the lack of foliar calcium response by white pins to soil conditions but data obtained by Scott suggest a response by this species, the tn well-drained Brown Podsolic soils giving rise to lower foliar 08 values than two less well-drained soils (see Table 3). Comparing the data of Plice and Gerloff it 3. with the above two, it appears that the range of white pine foliar calcium is from: about 0.3% to 0.7% with the upper range occurring in the freshly fallen foliage from high water table sites. (The calciun values reported by Alway _e_t _a_l_. are not included in this comparison since they seen out of line with presently existing data and were earlier considered abnormlly high by Plice.) A lack of response by sons pines is attested to by the data of Oviuten (1959) who states that Scots pine trees (associated with Podzols in Europe) show no foliar calcium increase with increasing availability of soil calcium on well-drained sites. Bard's study, Table 14, indicates the possibility that hemlock foliar calciun does not vary due to site differences between well-drained, nediun-tertured soils showing con- siderable variability in acidity, readily extractable calciu- and depth to carbonates. Most hardwoods in Bard's study contained higher foliar Cacontentswhengrowing onhighornedinnlinetillsas comparedtono line tills, but basswood and white ash mintained high and similar values on all sites. Scott's data, Table 3, however, show no foliar response by red nple within the range of site conditions studied, both values being lower than for the no line till site in Bard's study. The soil: in Scott's study were loam sands and sandy loam as compared to silt loans in tint of Bard's. h2 Foliar potassium response to site by pines is not apparent in the above studies although it has been shown that foliar potassium can be increased by fertilizing K-deficient stands of red pine (Heiberg and Ihite, 1951). In the case of sugar mp1s, yellow birch, eastern hemlock and ironwood, foliar potassium increases as the depth to carbonates in- creases. Ihite ash reacts exactly the opposite. I. foliar Iagnesiun response to site is apparent for white pins in Tables 3 and h. A foliar phosphorus response of white pins to site differences is apparent in Table 3. Most hardwood species show increasing foliar P with increasing depth to line, this relation being nest marked and consistent in the case of sugar naple and northern red oak. lite on foliar composition of ninor elenents in natural stands in eastern North America is scanty. Coaparisons between species on the same plots in Scott's study are liwited to the following: £322.22 22*...“ BEL—*1 m...“ Ihite Pine (Plot II) 275 325 550 Sugar Iaple (Plot II) 200 1.50 1000 Red Maple (Plot II) 200 150 1650 Ihite Pine (Plot III) 200 350 2650 Red Oak (Plot III) 150 150 h750 Caparisons between species on the sane sites in the study of Gerloff 33; g. are as follow: 1L3 Site No. Sagies m Fe PE Mn 76b Sugar Maple 157 805 Red Cal: 76 763 55 Ihite Pine 267 181; Red Pine 206 260 62 Balsam Fir 120 862 lhite Spruce 89 672 18b Ironwaod 239 28h8 Basswood 16h 121; 113 Butternut 196 1119 Black Cherry 159 620 6 Black Cherry 221 585 Black Oak 206 11459 White on: 126 1371: Red. (3k 125 1736 18a Iromrood. 278 968 Big-t. Aspen 106 So Ihite on: 101; 815 Comisons within species but between soil types in Scott's study are limited to the following: ‘ 31306100 8011 W ppm F0 ppm L1 ppm Mn Red mp1s , Bugabfifandy 200 150 700 Red mph Searbocro 133w 200 150 1650 . sand (mflszamy nor as 25 55 Ihite Pine Ridgebugsazwn 2 0 O haul-1 mu Pine Searboro 1mm 325 550 sand (11%)" 3 nor lhite Pine Kerri-c loam 175 325 2500 sand (BP‘) 3 nor white Pine Harri-c sandy 22S 35 525 loan (31*); nor "GH'é'Greledreuu-phic; HG=HuIicGley3BP=BrownPodzolic M; a chain of site-mate comparisons, disregarding leaf age, indicate that the foliages of ironwood, black oak, black cherry, butternut, sugar nple, red ‘ple and white pine contain higher concentrations of iron than those of white oak, red oak, big—toothed aspen and red pine. a chain of site-ate cmrieons indicates the following decreasing order of foliar 1m concentrations: red nple, ironwood, sugar mp1s and red eak > white pine, black oak, white oak and black cherry > butternut, basswood and big-toothed aspen. The latter three species apparently have especially low In contents. Scott's data indicate that white pine foliage contains higher concen- trations of alumina: than those of red oak, red nple and sugar ample. Evidence for foliar nngansse responses to site also exist in Scott's study. Rednplehasahigher foliar nnganese content onanorhunus- cweredflnnicoleyloawsandthnonamllhunus Greymdronorphicsandy lean. lhite pine he a much higher foliar naganese content on a nor hams-covered Brown Podsolic loam sand tun on less well-drained or finer-tortured soils. Orington (1956) found that Austrian pine and Douglas-fir both had about five tines as nuchfoliar Inganesewhengrswing onanacid sand as when growing on an alkaline sand. In twenty-two out of twenty-three foliar sasples involving sixteen species, foliar nnganese was present in litter quantities tun foliar P with greater amounts in the coniferous foliage thn in the hardened (families Pagaeeae and Betnlaceae only) foliage. Fron the above-national studies, it is clear that feliages from the sell-associated species, basswood, iromcod, button-mt, black cherry and white ash, consistently contain relatively high concentration of calcin and either relatively high or intemdiate concentrations of 1:5 potassiun and mgnesiun. The foliage of red oak, which is also associated with null humus types, seems to be outstanding with respect to its higher calcium content on high calcium soils and its higher I! and P content on low calcium soils. On both types of soil red oak foliage nintaine intermediate to high values of K and Mg and may contain relatively high concentrations of Mn on some sites. These latter chraeteristics may be related to the relatively high antacid bufferim capacity of red oak litter mentioned below. Species such as pines, hemlock and beech which are more often abundant on nor thn on null. humus types, have relatively low values of foliar Ca, P and Mg. White spruce and balsam fir, also comonly associated with nor huus types, contain intermediate values of foliar Ca uni P but thore is sons indication in the data that they usually have relatively low values of foliar Mg and occasiomlly very low values of foliar K. Species such as sugar neple and yellow birch which my be abundant on both humus types apparently have inter- ndiate and/or variable concentrations of nest of the elements studied so that the mture of the humus layer my depend largely on the svailable chemical elements and the associated species. Plice (1931;) studied the antacid buffering capacity ef the litter froa variou tree species on a range of sites near Ithaca, New York with the following results: h6 flies H-ion Inactivated 21 5g. of litter fine. out of I; added) Ell \ 3e963 3e8h Red on: Susir Mapl- Red Maple Iellew Birch White Ash Beech Ihite Pine lhite Spruce Balsaa Fir Red Pine Black Spruce Jack Pine Hemlock 3.1:; 3.1L 3.3-3.1: 3.2 3.0-3.1; 3.0; 3.0 2.h-3.o 2.0-3.0 2.0 2.0 2.0; 2.0 2.0 1.5 0.93 2.0 Plice concluded that both the chemical element content and the antacid buffering capacity were inflnential in determining the humus type. Polyphencls in Tree 3% a_nd_;_ litter Handlq’ (1951:), after extensive research, concluded that leaf pro- teins stabilised by mterials similar to tannins are an important factor in the processes leading to nor bums for-.tion. These stabilised pro- teins occur in the nesoplwll tissues and are, under certain conditions, so resistant to decomposition that the various parts of the debris in which they do not occur (especially the vascular tissue) decompose and leave, as a layer lying on the surface of the mineral soil, an amorphous residue of leaf neoplwll cell walls protected from decomposition by the h? resistant stabilized protein. Davies 32 2.3:. (1960) point out the likeli- hood that these taImin-like materials stabilizing leaf protein are P01313330”. Goulson gt 3. (1960), using paper chromtograply and electro- phoresis, unlined the polyphenols of fresh green leaves, dried leaves, litter and superficial humus from null-Brown Earth and mon-Podzol sites . They found a greater diversity and quantity of phenolic substances in the extract of the fresh European beech leaves from the nor-Podsol sites than in the extracts of European beech, sycamore and oak leaves from null-Brown Earth sites. There was a change in quantity of simple pclyphenols ranging from a maimm in living leaves, decreasing through senescent leaves to dead leaves to freshly fallen leaves, to a niniuun in decayed leaves and humus or stored dry leaves. However, tannin- stripping and hydrolysis-reduction of decayed leaves and superficial hulus released additional polvphenolic substances from both null and nor hm types. The above-antioned studies also point out that when polyphenols are poly'aerised beyond a certain molecular sise they are rendered incapable of am tanning action. Coulson gt :1. also state that polyurisation is favored by base-rich conditions and they suggest tint beech leaves falling on a base-rich surface lay explain m Europeanbeech gives risetoanullhunus typewhengrowing onbase- rich soils and to a nor humus type when growing on base-poor soils. The lower foliar calciua concentration exhibited by European beech on base-poor soils as compared to base-rich soils has been rcported by Handley (19510. It soon lihely that a low foliar base content as well as an acid soil surface would reduce polymrisation and thereby increase the taming potential of the pclyphenols which are present. Iv. I“ -9. '(‘I L. 11 A? ha as m: 2 en :92 he nuns n22 Ihile certain foliar characteristics my instigate humus type differences, it is probable that these differences are brought about by differences in the predominant groups of soil aninls attacking the foliage. lull appears to be the characteristic humus formd when earthwor- are the predominant group 3 transitional types of hu-swhenthedouinantsoilfaumarenillipedes, woodliceorlarvae cf the larger insects or termites; and typical nor when they are mites and springtails. The functional relationship see. to be that a greater proportion of the organic matter on lull humus sites passes through the alimntary caml of the larger soil aninls into the lineral soil (Russell, 1961). Although several studies have indicated extremely low rates of leaf tissue decomposition by micro-organis- when anilwals are con- pletely emcluded (v_:_l_de__ Edwards and Heath, 1963), evidence exists that niere-erganisu play a role in the preparation of leaf litter for use by the larger soil aninls and my be necessary for conplete decom- position of organic utter followim its ustication by the larger soil faun. Darwin, for instance, considered half-decayed leaves to bethe earthworms chiefarticle of dietandvanderDrift concluded froe his studies that the min result of the activities of nest soil faun is musical break-down of leaf litter (li_.de_ Handley, 1951;). Intypicalncrhulus types, the fungiere cfteneonsideredto be the predonimnt group of micro-organism, and they convert such of the leaf litter into their own protoplasnwhich is a fora that um nits. in particular can digest (Russell, 1961). A stun by Kendrick (1959) on Scots pine needles indicates tint successive waves of fungal colonisation initiates the deconpos ition process I. "J '11 1‘ ‘3' ft. 31' 1:9 and complete plus ical reduction is eventually brought about by soil fauna which results in an Oh horizon conposed largely of partly luneified aninl feces and numerous dead, demtiaceous hyphse, conidio- phores and conidia. A similar process my lave led to the observations of Ewell (1935) who stated that heavy greasy nor humus layers in the northeastern United States sea to be built up chiefly by dead fragments of brown lyphee. In lull hams types, however, bacteria are probably the nest is- W Iicrobial agents of decomposition (Russell, 1961) and they seem to be acre abundant in the presence of earthwor- (Went, 1963). Several conditions seen to be related to these differences in soil. organic: populations and activity. Aside from moisture requirenents (Lids; Hilde, 1958), studies show that earthworm seen to have definite preferences for the leaves of certain species of plants. One such stuw in Europe indicated tint in general, the earthwor- character- istic of ml]. soils shows preference for the litter of elnand birch, consuls onlysnllasounts of beechandoaklitteramldonotconsme pine and spruce needles at .11 (119.; Handler. 1951;). In the United States, they show similar preferences for foliage rich in bases (such as ash leaves) a reluctant ability to kindle the tough leathery leaves of oak and beech, and a distaste for acid conifer needles (Spurr, 19610. Edlards ani Heath (1963) noted that tanned European beech leaves wereeot eatenbya soilfaum including earthernwhilegreenor yellce leaves of the ease species were heavily attacked. Recent finding. an. indicate that m1 can utilise organic substances containing concentration of phenols sufficient to inhibit bacterial 50 attack (Kononova, 1961 and Basaraba, 1961;). A related study indicated that nrhsd differences in faum populatiom between litter samples rich in neelia and salples not so endowed were correlated with the lower pH an! higher tannin content of the melia-rich samples (Kuehnelt, 1963). Processes 2d Predicts 2f _th_e_ m Horisons Iafonl (19h?) neasured the oxidation-reduction potential of finely packed null ani nor nterial which Ind been allowed to stand overnight in a water-legged condition. He found that as a rule mll humus has a positive oxidation-reduction potential whereas nor has a very low negative potential. Other studies indicate tint ferrous iron become prelimnt at Eh (endation-reduction potential) values below about 0.2 volt during periods of intense microbiological action. We is effected sinilarly but aluminum is not (Alennder, 1961). In the Podsol Region of lishign, McKenzie at g._l_.. (1960) found that the A1 (or Vb) horizon of a well-drained sandy Podsol (Kalkaska sand) ahihlted the lowest redo: potential inthe profile andwas consistently at its lowest seasonal value in winter under a cover of snow during the two-year duration of the study. Bonell (1935) states tint in the latter part of the 19th century it was noticed that null and nor give extracts of different colors, either with distilledwater or with weak anonia. He claim that mll extracts have a "less inteme huaus color." This water soluble, intensely hams-colored extract from nor is probably similar to the extracts obtained by Berselius who is credited with the original description of crenic acid and apocrenic acid (Konenova, 1961). According to Kononova, apocrenic acid is formed by the atmospheric oxidation of crenic acid. She further states that Berselius describes 51 crenic acid as having a yellowish color, a sharp taste and being elerphous while apocrenic acid is described as having a brownish color; both possess acid properties. Koneneve also mentions that Berselius nde couprehensive studies of the K, Na, nah, Ba, ca, lg, n, In, F.“ and F.” salts of crenic and apocrenic acids . Ac- cording to a nnsber of Russian soil scientists, erenic acid is profusely produced by the activity of fungi in Podzol humus hyers (Vii-uni, 195?; fillians, 1911:). Oden (vi—dz Kononova, 1961) introduced the tern 'fulvie acids. for the group of hueic substances occurring in post waters. He described these substances as compounds of high nolecular weight chracterised by a reduced (lees tlnn 55%) carbon content and high solubility in nter, alcohol and slhli; their salts are also readily soluble in water. At low concentrations these substances are slightly yellow in color. Il'hese chancteristics apparently led Oden to the aseunption tint fulvic acids are amlogous to crenic acid and apocrenic acid. Subsequently, fulvic acid was studied try new investigators as the acid salable portion of alien-extracted orpnic setter (Konenova, 1.951). These studies indicated the presence of pentceans, uronic anlvdride, asino—nibeges, phosphorus, several sugars, and phenolic glucoeides. Infra-red spectroscopy and I-rey analysis revealed that fnlvie acids possess 'etructsral units“ of srontic compounds, nitrogen- containing substances and reducing substances. Recent studies by Wright and Schnitser (1963) on s Camdian Podsol indicated tint the extracted organic utter fros the Oh and Bh (Ihbi?) horisons contained 30% and 85% mm acid, respectively. in mob n 60%of this fulvic acidwae conposed offunctionalgroups suchas 52 carbalyl, hydroxyl and carboxwl which appeared to be attached to a predominantly sronntic “nucleus ." Iarkov (1951;) denonstrated that the nobility of complex organe- nineral eta-pounds of fulvic acids with R203 depends on the oxidation- rednction coalition of the soil nediun produced by the seasoml noieture rcgin (Lid: Kononova, 1961). else-triad (1957), Kaurichev gt :1. (1958) and Goulson .932 g_l_.. (1960) untion that polyphenols capable of forming complexes with iron are present in Podsol humus layers. Bloomfield considered the polyphenels to be inportant in the dissolving and reduction of R203. Goulson at .‘l’ showed that D- and epi-catechin (two of the njor polyphenels of beech leaves) are capable of reducing iron and obtained evidence of the fornstion of ferrous iron-polyphenol complexes. Aeration had an adverse effect on the forsntion of these complexes. The nevenent of sluninun, however, did not appear to be influenced by these polyphenols but was nore effectively leached by strongly acid attractants, the nest effective of these having a pH of 3.72. Hesselnn (1917) concluded fron his studies that, in general, conifer nor hunus is! characterised by active amonification but no nitrification. Resell (1931) found that intensity of nitrification in nor lnmus layers was correlated directly with pH. Moon and Sowden (1959) stated that their coniferous litter-Podsol soil (horison not nentioned, presunbly the A heriaonswwere used) nixture produced no nitrate during the course of their erperinent (165 days). Lent (1932), working with New England forest soils, found tint sulltypes ofhu-is foundinfast-growingtnrdweodstands nitrified ,‘ 53 to a considerable degree with the accumulation of only a relatively can alount of anonia. 0n the other hand, large quantities of a-onia accumulated in the thick nor bums found in nture hemlock- hardIeod and nture white pine stands. Studies of the possible causes of this pattern revealed that pH, Ge content and sometimes total N were correlated positively with nitrogen transformation. These correlations were quite pronounced in the Oh horizons. The addition of line generally caused the forntion of nitrates at the expense of ammonia, though no appreciable effect could be obtained in the humus layers from red pine plantations. The work of Lunt agrees with earlier studies by Noses (1930) and concurrent research by Renesov (1937) who both found tint nitrification is greatly retarded in Inmus layers under stands of spruce and pine. Anonification, however, took place readily and accumulation of large alounts of anonia occurred under these conifers. These researchers explained the lack of nitrification by their discovery of a negative correlation between the bitunin" content and the rate of nitrification in the humus layers. With bitunin contents of 5% or sore nitrification was practically nil. Ghee and Baker (1951;), working with Camdian Pedsols, found that heavy applications of calciun carbonate were required before an of the added anoniun nitrogen was converted to nitrate. On an acid Canadian Podsol under nple, Corina (1958) found that linestone- phosphte fertilisation had greatly increased the number of an- unifying and nitrifyim bacteria 8 years after treat-eat; ’Bitusim is apparently a collective term for humus constituents such as fats, waxes and resins which are soluble in alcohol and benzene (see Vilemkii, 1957). 5h simltaneously, an increase in ammonifying :31; occurred under conifers following this type of treatnnt. When amoniun sulfate was perfused through the 10(0) horizon from the maple plots which had been treated with lisestom and limestone phosphate, a fairly rapid oxidation of NHh-N to NOB-N took place. In order to obtain similar nitrification rates from the unleaded plots, large amounts of M03 had to be added prior to incubation. No mention was made, however, as to whether or not the coniferous 0h horizon could be stinmlated to form nitrates. Renescv (1937) states that the accumlation of large quantities of a-oniun in humus layers my promote dispersion of some of the hums and thereby convert it into forms nore liable to leaching. CHAPTER h. TIME RELATIONSHIPS In his monograph on soil studies in the region of coniferous forest in northern Swoden, Tam (_v_i_d_e Jemw, 191d) states that in a drained lakebed perceptible podzolisation can occur in 100 years ; under a mattress of raw hums, enough Podzol formtion had taken place during that time to permit a photographic recording of a thin bleached A2 horizon and a dark orterde sons (Podsol "3”). A study in Alaska made by Cracker and Dickson (1957) indicated that in 200 years of soil development, a trace of Podzol formtion was evident in the sandier materials although no profile descriptions nor chemical analyses were presented for corroboration of the visual evidence. The youngest well-developed Podzols described in this area were 3000 to hOOO years old (Chandler, 19142; Crocher and Diclmon, 1957). LEEELfiW mam—hires land surfaces in the Podzol Region of Michigan have been exposed for periods of tine not exceeding 13,000 years (Zusberge and Potsger, 1956). For areas covered by the Valders substage of Wiscomimn glaciation, this min lisit is reduced. According to the nest recent radiocarbon dates (Broscker and Farrand, 1963), these areas have been exposed for a nximn of 11,850 years. In the Lower Peninsula, Valders drift is alaost confined to Podsol Zone III (1133 Zunberge and Potzger, 1956). In Cheboygan County, Michigan (Podsol Zone III), Fransneier (1962) studied a chronosequence of Podsols foned in sand primry materials. This sequence consisted of weakly developed Podsols recently under pre- doaimntly pine and oak on lake ngon and lab Nipissing surfaces, a 55 {I (t 56 acre strongly deve10ped Podzol which recently supported mostly hemlock or balsam fir on a lake Algonquin surface and somewhat more strongly developed Podsols (on the basis of darker upper illuvial horizons) under northern mmoods (one site) and aspen (another site) on Valdsrs noraines. 1113 E2 m _t_g Regional Changes in Climate and Vegetation If the regional climate and vegetation had been uniform during the last 13,000 years, the time of land surface exposure would be the only variable soil-forming factor in the Podzol Region (providing topograplw and parent mterial are held constant). Since the Podsol Region of Michigan he been subjected to post-glacial climte and vegetation changes, land surfaces of widely different ages must have been affected by different combinations of climatic factors. For instance, Isle Royals was coupletely inundated until post- sub-Duluth tine (Hough, 1958). Potzger (1951;) found that a bog on a post-sub-Duluth, pre-iiinong surface 900 feet above sea level indicated an initial post-inundation forest of pine and spruce. Two bogs on a pcet-Minong, pre-Nipissing surface 650 feet above sea level indicated an initial post-imndation forest dominated by pine. The bogs at lower elevation, which are all on Nipissing and post-Nipissing surfaces, indicated that initial post-Nipissing forests were damn-ted by pine, spruce and birch. Potsger states tht the pine-donimted parts of the pollen profiles undoubtedly represent the "flier zerothersic' period in that area; thus initial soil fomtim on the post-linong, pre-Nipissing surface took place during a different clinte-Ivegetation regime than existed during initial soil foamtion on either the older or the youmer surfaces. 57 The Isle Royals pollen data suggest that pine forests were re- placing spruce forests by sub-Duluth time (around 10,000 years B.P., Li_de_ Broecker and Fan-and, 1963) and were prevalent in early post- }linong ti. (correlative with lake Chippewa times in the lake Michigan basis according to Bough). The tine range of 10,180 1: 160 to 9,150 I 130 years B.P. given by Fries (1962) for the decline of spruce in nearby laks County, Mimesota seem to be compatible with the lake chronology of Breecher and Farrand (which entails a drop from lake Algonquin levels to lake Chippewa levels between about 10,500 years ago and 9,570 1' 150 years ago). The data presented by Rios suggests that pine was dolinant from 9150 2 130 years B.P. until the late post-glacial increase of spruce and fir occurred. His data also suggest tint Jack pine was the predonimnt pine until 7300 1' 1140 years B.P. Based on the above studies, that of Potzger (19h6) and those of Iilson and Webster (191m and 19h2b), the following post-Valders clintic and vegetatioml conditions .are inferred for the Upper Peninsula of Michigan: Tin (_B_l_’) Cli-te figgioml Forest Yam 11,850 I 100 to Cold, nesic Spruce-Fir 10,180 r 160 10,1801'160te Warner, less mic Increase inpineandtherso- 9,150 1'. 130 philous deciduous species; hemlock appears in some areas 9,150 I 130 to Even less nesio Pine dominance (ninly Jack 7,300 2'. 11:0 pine); low or no hemlock; low white spruce; 1w thernophilous deciduous species 7300:1110“ More sesis Increase inwhite pins; in- approx. 3500 crease or reappearance of henlock; increase in birch approx. 3500 to Cooler and even Increase in hemlock, spruce lumering are more nssie fir and birch 58 The following are estimates of the same types of relationships for the sandy prism-y nterials of northern Loser Michigan: 2'33 (2.3.) 13,000 11,000 8,500 7,000 3,500 2,500 Climte* 0001 to cold, noist 0001 to cold, noist Moderating clinte Farming climte Warmest and driest since retreat of ice Deterioration (cooler) Regional Forest Vegtation'“ Spruce-Fir Spruce-Fir Spruce, fir and Jack pine Pines, oak Pines, oak and northern hardwaods Pines, northern hardwoods and oak i'Based on Disberge and Potsger's interpretations “Based on Kilburn's work In general, both sets of data suggest tint the older sandy land surfaces in the Podsol Region of Michigan were initially exposed to a cool to cold noist clinte and supported pioneer stands of spruce and fir. The niddle-aged surfaces were initially exposed to a warmer and drier clinte and were probably initially forested by such species as pine and/or oak on the drier sites with arborvitae (white-cedar), helsen fir and white spruce on the cooler and more noist sites. Surfaces expend since lake Nipissing or lake neon tines were initially exposed to a cooler and possibly noister clinte and probably supported pioneer forests with less oak tun occurred in the initial forests on the middle-aged surfaces . Aspens also my have been involved in the initial stages of early, middle and late successions even though their fossil record is hissing (Lid: Wright, 1961;). Sill]. percentages of nple, hemlock, beech and other non-boreal 59 hardwoods appear at the end of the spruce-fir pollen zones in peat profiles fro. northern Lower Michigan and north central Wisconsin. Indication of beech-uple forests in these areas, however, are restricted to the last 3000 to 14000 years. Another tin-related factor which s hould not be overlooked is the tine during which the “lake effects" have been in existence. These phone-en are directly or indirectly responsible for web of the clnraeter of the climate in the Podsol Region of Michigan. One of the nest striking of these‘ effects is the fall and winter precipi- tation. For steeple, Grand Marais, Minnesota, receives about 63 inches of snow per year while Houghton, Michigan, receives about 120. Male are characteristic in the Houghton area but not in the Grand Innis, Minnesota area. Milwaukee, Iisconsin,receives about 39 inches of snow per year while Grand Rapids, Michigan, receives about 69. Podaols are present only in the latter location. ..Inwinter, the warner the lake waters are, the greater will be the instability of the air which has passed over then and the greater will be the snow- fall on the downwind side of the lakes . Consequently, relatively little snowfalls onthe downindside ofthe lakeswhentheyare from ever. By the cal of February, 1963, ice covered 95 per cent or lore of lakes Superior, Michiganand Huron. lalne Michiganwas from over north of a line between Millaulnee, Wisconsin, and Mashegon, Michigan, fru February 26 to the end of the nonth, the only other such kncsn occurrence having been in February, 1936 (Heather Bureau, 0.8. Department of Gomerce, 1963!). Every section of the state received less than nor-s1 precipitation in February, 1963 andtheUpperPeninsulareceivedless thannonalinMarch, l963as 60 well, with departures from the normal being greatest in the western part. Chatta- arxl Munising (both in Alger County, Michigan) had negative precipitation departures of more than one inch in March. The following table compiled fron Climatological Data (Weather Bureau, 11.3. Departnent of Connerce, 1963b, 0, d, e, f and g) illustrates these 1963 departures (in imhes) from norml precipi- tation in regions at comparable latitudes but varying in their clinatic dependency on the Great lakes. East Central Northwest West Upper Minnesota Wisconsin Mic%n February, 1963 £55 3.5 ”11, MB 'I‘OeOh "Oelh .1e61 Southeast East Central West Central Minnesota Wisconsin Lower new February, 1963 20.5 5.33 . ”h, MB «00.08 40.50 40.50 The above data suggest that during nest of post-Valders—pre- Nipissim tines lake effects nust have been less pronounced the at present if it is inferred that: (l) winters were somewhat colder during the spruce-fir period and (2) that a drop of lake levels to ex'trusely low levels took place during the decline of the spruce-fir period. Durim the lake Chippewa period, spring and smea- temperatures in southern Michigan would Inve logically increased whereas fall and winter tenperatures would likely have been more continental, sinilar to those in southern Wisconsin and northern Illinois at that ties. Fall and winter precipitation would have been co-ensurately lower aswell. IhilethebulkofthePrairiePeninsulanyhavebeenfornd prior to this period, as suggested by Benninghoff (1961;), the lake Chippewa pericdseen tcbea likelytine forthefecntionofthe soutmrn Michigan prairies with the disjunct Newaygo prairies developing 61 as a partial result of: (1) their position at a latitude betw9en that of Southern lab Chippewa and lake Chippewa (see Hough, 1958) and (2) the prevalence of westerly winds (Weather Bureau, 1959). The subse- quent rise of water to the lake Nipissing level must have initiated an increase in the ”lake effects“ which would mean cooler springs and ms, less annual evapotrenspiretion, milder fall and winters and more fall and winter precipitation. These changes plus a regionally cooler clinte beginning between 2000-2500 years B.P. (Deovey and Flint, 1957) could possibly explain the increase in white pine indi- cated in the upper levels of several pollen profiles in the laer Penimula of Michigan (post-Nipissing phenomenon according to Zunberge and Potsger, 1956). These climtic changes could also have aiml- tansously favored the increase of: (l) mesophytic species such as henleck and mrthern hardIoods all (2) the rate of leaching all! Podzol development. Thus the lupothesis (Zunherge and Potsger, 1956) that lake Wipissing tines were the nest xeric in Michigan seen untemble unless it is assuned that expansion of beech and hemlock can take place under such conditions and a rather drastic regional dryness occurred din-in that period thereby overeating the lake effect. 113: 2th m 33 32°.“— Chanes _i_n Soil-Forming Factors The energence of sons land surfaces mt have been rather rapid while others surged slowly (Veatch, 19140). The slow subsidence of lake levels could not only affect the natural drainage of developing soils but promisity to the lake could keep sumer temperatures relatively low and winter temperatures relatively high compared with more inland positions. For maple, if lake Nipissing receded 62 steadily down to the lake Algona level on a stable land surface, the water table in lake Nipissing beach sands would have been within 5 feet of the soil surface for approximtely 500 years. If lake Algona levels dropped steadily to those of the present lakes, this time would be about 1250 years for lake Algona beach sends. it the other extrene, deep out-ash plains subsequently dissected by saltwater from the retreating ice front were high and dry shortly after their depo- sition. Surfaces exposed by the drop of water levels from lake Algonquin to lake Fayette at an average rate of at least 1!; feet per 100 years (& Bough, 1958 and Breecker and Fan-and, 1953) were soon without lacustrine water table influence as well. CHAPTER 5. PODZOL PROPERTIES AND PEDOGENIC PROCESSES 32. m constituents Determinations of total Fe203 in Podzol profiles usually indicate that at least one illuvial (I) horizon has a concentration twice as great as that of the e‘luvial (E) horizon and that the eluvial horizon has at least 20% less than the 00’ or I!) horizon (v_i_d_g Lunt, 19323 Iilde 33 31,, 1910; Wicklund and Whiteside, 1959; Muir, 1961). These studies also indicate that I/E* ratios of total A1203 are lower than those of Fe203 except under pure stands of conifers nhere the reverse is true. muvial peaks of total P205 in the Podzols studied were incompicuous except where crtstein was present. Otherwise, P205 unnu- occurred in the hams layers. In the Wisconsin Podzols, total K20 concentration were considerably lower in the eluvial horizons than in the other horizons. The New Brunswick Podzols, however, did not exhibit this type of distribution pattern for x20; instead, the values were rather constant down to the lower I or upper I horizons where they increased. Total CaO concentrations in the Podzols stmiied were highest in the 0 horizons (exclusive of profiles having carbonates in the P horizons) and lowest in the E horizons. hummus concentrations of MgO were present in the E horizons with nui- in either the I horizons or the P horizons of carbonate-free profiles. Ihere total 8102 was deternned in the above-mentioned studies, the results indicated that Podzols of all textures had elnvial horizons with 8102 concentratiom above 79$. In the lisccnsin sands, Podsol development seems to increase with increasing l/P ratios of total 8102. in increase of this ratio in the HewBrmwickPodzols is coincidentwithagreater net increase in 11-1/3 gilluvial horizon s zone 63 6h illuvial 11203. Iasseglou and Ihiteside (1960) determined the amount of soluble (Whole extractable) aluminum in some Michigan Podzols containing fragipam and found that the morphological degree of Podzol deve10pnent varied directly with the content of soluble, illuvial A1203. Franneier (1962) (1er the citrate-dithionite extractable iron and aluminum of several sandy Michigan Podzels. Concentrations of the sesquioridss increased with the morphological degree of Podzol development (based on color of the upper illuvial horizon) up to the moderately developed Blue lake soil. This moderately developed Podzol (a continuous ortstein is not present) with the greatest concentration oil-.203 of am soil in the studywas formed inValders-aged parent nterial. Free the available evidence, this soil supported a conifer (white pine and henleck)- northern hardened stand prior to beirg clean- cnt. The highest concentration of £1203 was shared by the above-mentioned Pedml and an Algonquin-aged Pedzel which supported a pro-disturbance stand of moplutic conifers (either hemlock or fir or both) with somewhat of an adnixture of red nple (A_ee_r_ w). The above study further indicated that in the lowest illuvial horizon of one of the weakly (Rubicon series) and all of the moderately developed Pedzels (Kalkash and Blue lake series) extractable aluwinm concentrations were consistently higher than those of iron. In these profiles, the upper illuvial horizons consistently contend the highest concentrations of translocated humus. In addition, these horizons had tb maxim R203 concentration in each profile except where ortstein chunks were present. Available phosphorus distributions in Franneier's Podzels are also 65 of interest. In the weakly developed Pedzols, the illuvial zones show urinal concentrations in their upper horizons while in the Pedzol sequue of the moderately developed Podzols, mdnun concentrations of available phosphorus occur in the lower illuvial horizons. This distribution in the moderately developed Podzols is nest obvious in the Blue lake profiles which have Bhir (lhbi) horizon concentrations that are alweet twice as high as these in the overlying Bh (lhib) horizons. Total analysis of one Blue lake profile revealed that the upper illuvial horizon had actually gained in total anounts of phosphoru. The next illuvial horizondownportrsyeds net loss of total. phosphorus but contained almost twice as much available phosphorus as the horizon above. The 953113 Constituent. 93 Hanged Mechanism 9; Eluviation £1 E Illuviation _i_n Pedzels Pedzel illuvial horizons are characterized by accumulation of sesquiddes 3 however, organic utter accmulations are invariably present as well. Franz-sier's studies indicate that organic natter: extractable sesquionlde ratios are consistently greater than unity in these horizons regardless of the degree of Podzol develop-ant. If the extractable sesquioctides represent that portion of the total which is ecving or has moved, and the amputation of organs utter content is realistic, then the data indicate an essential role for organic wetter beyond the nere creation of acidity. Because of the canipresence of organs utter in Pcdzel illuvial horizons, nest of tln modern lupotheees consulting the nehanisn or nectarnsm of iron and alunimn wovemnt into the illuvial horizon involve the leaching of organic substances (from 0 horizons or live foliage) which are capable of (0 66 combining with iron and aluminum (Stobbe and Wright, 1959). Several Russian investigators Lug Kononova, 1961 and Vilenslcii, 1957) believe that crenic acid is responsible for: (l) the dissolution of calciul carbomte, (2) reaction with iron and manganese compounds and (3) disruption of holidte thereby releasing silica and alulinmn, the latter foe-ling a create (i.e., a salt of crenic acid and aluminum) which is water soluble. Upon reaching a zone of oxidation, these cremtes of Fe, Mn and A1 are converted to apocrenates which are not nter-eoluble and thus precipitate out of solution (see Kononova, 1961 and Vilsnskii, 1958). lright and Schnitzer (1963) postulate that the iteration of fulvic acid night be visualized as arising through sale alteration of humic acid, including an increase in oxygen content, an increase in carbonyl groups at the expense of aliphatic and/or alicyclic laterial, and a decrease in carbon, ludrogen and nitrogen. With increasing oxidation the nterial becones more water-soluble and eventmlly dissolves in water. is the dissolved fulvic acid nerves down the profile it conbines with pclyvalent cation such as iron and alulmue to for. water-soluble cenplexes sole of which probably involve two or more donor groups of the ligand resulting in the formation of natal chelates. The authors suggest that there is a strong possibility that on its pith down the profile fulvic acid form, at first, water-soluble nultidentate chelates which later my precipitate lower in the profile upon reacting with acre of the es. netals orwith extremelysnllaeounts of ionic calcimand/or Hen-sin. Wright and Schnitzer also suggest that F's-organic ntter conplues are more susceptible to flocculation by Ca and Hg than the Al- organic fitter comlms which any result in a deeper penetration of the latter. 67 On the other hand, certain researchers believe that organic acids such as oxalic and citric lay fora conplexes with Fe and Al. which subsequently nove into the illuvial horizons (see Stobbe and Wright, 1959). Recently, the importance of polyphenols has been stressed in conjunction with the mnt of iron (Bloomfield, 19573 Goulson, _e_t 91., 1960; Davies _e_‘_t_ g... 1960). Coulson at 3;. treated nodel soils (con- sisting of slain and diatmaceous earth inpregmted with ferric chloride) with catechin" solutions and fresh European beech leaf extracts. Both treatments produced dark-colored subsoil bands in which substantial lacunts of ferric iron had been converted to ferrous iron. Extracts of greenbeech leaves frona nor hums sitewere acre effective than those from a null. huaus site. Siailar treat-cuts of Triassic sand calms also resulted in the reduction of iron but apparently no subsoil bands were produced; the ferrom iron appeared to be present in a complex fora possessing no residml electric charge. Iaachingofalmm seened tobe related onlytothe pH ofthe solutions. Regarding the Podzol-foraing process, Davies 33 31. state tht, upon reaching the soil, the fate of polyphenols is deter-ined by the soil reaction—the acre acid the soil, the acre stable the poly- phenel. Further, these investigators suggest that the polyphencls responsible for movement of iron in soils are likely to be these washed free the growing leaves into the soil, and not those fro. litter or hulls. Therefore, if the soil. is acid (pH of 14-5 is optima), the leaf polyphenols will. readily reduce ferric iron and fun stable *Catechins are a group of polyphenols found in tree foliage. f‘ 68 conplaxes with the resultant ferrous iron. Since these complexes are water-soluble and non-ionic, they will nova freely in the profile until they are deposited in a clearly defimd horizon, thus forming the Podzol illuvial zone. These authors do not postulate a mechanism for this deposition, however. Concerning deposition in the illuvial horizon, several workers (see Stobbe ad Fright, 1959) have stressed the importance of oxidizing conditions and nicrobial attack of the organic utter as it moves into this zone. Bloomfield (1957) suggested that drying and/or aeration my brim about the precipitation of sesquotzides. He also found that the mobilization of sesthimdes is associated with the sorption of the eta-planes on the aineral soil particles, particularly on the «equi- azides. firtin (1960) concluded, based on his studies of the illuvial process of Podzols, that the sisultaneous presence of L1, Fe and hunts in the illuvial horizon can be accounted for solely by the flocculating properties of Al ions. m 93 20% Develop-ant in Michigan Most studies of Podzcl genesis in Michigan have been of a moral- ogieal mture ani this subject is thoroughly reviewed by Frenzneier (1962). His study of Podzol sands led his to postulate the following course of developuent. An early accululation of available phosphorus occurs in the very slightly developed illuvial horizon along with couperatively low concentrations of iron and alt-1mm. This stage is referred to as the W phase." Following this stage, an "organic-accumulation phi-0" bogim and n: horizons fora. euquoandoe and probably silicate clays continue to be nobilized in this phase, but accumlations of these 69 constituents tend to be in different horizom. Several mechanisms of nobilization and various combinations of the active components are probably operative during this phase. The active sesquioxide and organic cemenents are adsorbed or precipitated as amorphous coatings on slightly crystalline coatings in the Podzol illuvial horizons developed during the I'inorganic phase.” The thickness of the amorphous coatings gradually imreases until they flake off and bacon inter- granular deposits. Here, acting as nuclei for further precipitation and adsorption of laterial from solution, they cause an increase in the amount of intengranular material. Sims these aggregates are relatively weakly held together, chemical, physical and biological agents prevent than from growing indefinitely. Host of the aggregates are about 0.02 to 0.1 In in diameter. is the large pores become filled with this debris, the capillary pore space, readily available water- holding capacity, exchange capacity and enchngeable bases increase. Condition are thus nde acre mesophytic and these changes are associated with the maple-beech succession of the pine-hardwood association. 0!: the basis of his chronosequence stuck, F'ranzmeier further concludes that, during the entire course of Podzol forntion, musical weathering caused a breakdown of sand grains to silt size (especially near the soil surface), and the total clay content of the solun increased. 4' CW 6. WIRED SOILS DEVELOPED IN IELIDW‘BH SANDS IN THE PWZOI. REGION OF MICHIGAN The Podsol Region of Michigan contains a very large acreage of soils developed in yellowish (Munsell hues of 7.5m to 101R) sand. However, compared to Gray-Brown Podzolic Region soils developed in yellowish sand, those in the Pcdsol Region vary greatly in their norphclog. V Iell-dnined Podzol Region sand soils, exclusive of those on the younger land surfaces, may belong to any one of the following three great soil groups: (1) Podsol, (2) Brown Podzolic or (3) Brunisea. In addition, transitions betueen (1) and (2) and (3) my occur. However, thsBruniseaintergrade onlyoccurs inPodsol Zone I (Figure 2) under white pine—mixed oak (ninly white and black) stands adjacent to the Brunlses areas (personal observations by the writer). On the younger, well-drained land surfaces Regosols can also be found (persoml obser- vations). The plusical geography of the well-drained sand soils in the Podzol Regina is presented in the following sections. mammals Restart Lnd This very weakly developed Podsel is found only on surfaces abandoned by Iahe ngm (personal observations arsi Veatch, 1953) near Labs Michigan or labs Huron. It has a shallow (usully less than 2% feet deep) solua, contains no dark (Munsell values and chrome less than h/h) illuvial horizon, nor reddish horisons (hues redder than 1013), and overlies a calcareous or alkaline G horizon. In Delta County under relatively un- disturbed conditions, the hunus type is usually a strongly acid to very 70 a, strongly acid nor or duff-null. The mtural vegetation (recent) on this soil in Delta County varies free an overstory of white and red pines with an understory of arborvitae (1113a occidental-ls) and balsam fir to stands of northern hardwoods, red oak and white pine. According to personal comunication with S. G. Shetron (formerly of the 8.0.5. new graduate student at the University of Michigan), the undisturbed Eastport sends in the northwestern part of the Lower Peninsula have similar vegetation types. The Sanilac County (east central Lower Peninsula) soil survey report (1961) indicates that “scattered scrub oaks" may have been a part of the natural vegetation on Eastport sand in that county. Eastport sand has developed under a more marine clients than exists in areas further fro: the Great lakes. This is particularly true on the west coast of Lower Michigan where mean January tomeratures my be as much as 5°!" warmer than more inland stations at the sans latitude. Mean July tenperatures are only slightly cooler in the coastal areas, however. The frost-free season say be as much as a month longer on the coast than inland. Eastpcrt sands which have developed in Podsol Zone III receive apmciably more fall precipitation and armual snowfall than the east coast Eastpcrt sands (11.8. Weather Bureau, 1955; Brunnschweiler, 1962). no.2 are 22 Deer Park sand, like Eastport sand, occurs on lake ngon surfaces. It, too, is a veryweakly developed Podao1(ne dark illuvial horizon and no reddish illuvial horizons), but the solun overlies several feet of acid sand. Generally, this soil has a deeper water table than Eastport sand sites (Veatch, 33 3., 1929; see Appendix v). l"*l‘ 72 The natural vegetation on this soil is quite uniform, being composed of red, white and jack pines with some admixture of northern red oak (Quercue rubra and Quemus rubra 1° borealis). noises 22 Rubicon sand is a weakly developed Podzol; i.e., it has reddish illuvial horisom, but no appreciable (thicloer than an inch) dark horizon in the illuvial part of the solum. The solum is variable in thickness, but the mxieum my be greater tlmn either of the two preceding soils, tongues sometimes approaching 5 feet in Delta County. This soil can be found throughout Podzol Zones II and III and my occur on ary land surface that is 3500 years old or older (persoml observations by the writer in Delta and Alger counties; and Veatch, 1953). The natural vegetation of Rubicon can! (as defined by the Natioml Cooperative Soil Survey) is quite uniform in the Upper Peninsula, being composed of red and white pines minly. Northern red oak and red mple my be present in emll quantities in undisturbed stands, however. In the lower Peninsula, northern red oak was present in most stands and probably in greater numbers. In addition, white oak was sometime present (Stewart, 1927a and 1927b and Elliot, 1953). In Delta County, white pine is dmimnt and hemlock is present where Rubicon sand grades into the moderately well-drained Croswell sand. Where the imperfectly drained Au Gres sand is encountered, balsam fir is frequently a component of the forest as well as hemlock. As the degree of Podsol development approaches that of a moderately developed Podsol, hemlock and northern hardwoods my both be present; in this case, hemlock is usually more prevalent than northern hardwoods. In Kalkasln County (Podsol Zone III 73 in the Lower PardnquA), Stewart (1927) describes a virgin timber stand on Rubicon sand which consisted of basal area percentages of: 69% white pins, 7% red pins, 20% hemlock and 14% northern hardwoods (sugar mple, beech and yellow birch). This stami data suggests a transition to the moderately developed Podzol, Kalkaska sand. A more typical maple is the basal area composition of another virgin stand in the same county which contained 23% white pins, 75% red pine and 2% hardwoods (red mp1s , red oak and poplar). Stewart described a third virgin stand in the adjoining county (Crawford) which consisted of 100% red pine. bunch and Kalhsh sand is a moderately develmd Podzol; i.e., the illuvial zone contains both dark horizons and reddish horizons, with the dark horizon averaging several inches in thickness. In addition, at least one dark horizon nust be continuous and an inch or more thick. Ialkasln sand is largely restricted to Pcdzol Zone III; however, it also occurs near Lake Huron in Podzol Zone II (Schneider, 1961). It is more frequently found on surfaces which pro-date Lake Nipissing but can be four! on Ialne Nipissingqabandoned surfaces in Delta County*. The recent natural vegetation of Kalkash sand varies from nearly pure stems of white pine (observations by the writer and Sanilac County Soil Survey Report) to 100% northern hardwoods. m Alger County, one virgin stand contained 711% sugar mple, 18% beech, 7% yellow birch and 1% iromood (933m 3%) while another contained 115% beach, 28% supr mple, 22% yellow birch, 11% red mple and 1% balsam fir 'nlhsh sand on Lake Nipissing surfaces: (1) ssl 5:} Sec. h r to N n 21 11, Rapid River Quadrangle, use: Geological Survey; 1958, (2) MM». 23 T 39 u n 22 w Rapid River Quadrangle, USDI Geological Survey; (3) saunas». In T to N R 20 v Garden Quadlengle, USDI Geological Survey, 1958. 7h (Stewart, 1929). Other stands (non-virgin) on Kalkaska sand in the same area contained black cherry (Pr—ulna; serotim) and hemlock in addition to beech, birch and naple. The nearly virgin Cross Village stand in Ell-at County (Lower Peninsula) is a mixture of northern red oak, hemlock, sugar mple, beech, white pine, hornbean and yellow birch (personal comication fron J. E. Cantlon, Botany Department, Michigan State Ultlversity). sue: 22d; Wallace sand represents the maxim development of Podzcl sorphology in fichigan. The bleached eluvial horizon is irregularly thicker than those of other Podsol sand soils; the illuvial zone is highly indurated (contains Inch ortstein) in addition to having the dark and reddish horizon. Eluvzlal and illuvial tongues are characteristic, the latter sonstins extending to depths of 5 feet. Illace sand occurs locally throughout Podzol Zone III and to a lesser extent in Podzol Zone II where it is not extensive enough to be upped as a single unit but it is cabined with associated soils such as Weare fine sand (Johnsgard, 1950. Wears is now correlated with the Kalhska and Rousseau series.) and Ballasts sand (Schneider, 1961). In Delta County, Iallace sand can be found on an land surface which pre-dates Lake Algom (persoml observations)*. In nest soil survey reports (Voatch 93'. 31., 1932 and 1931;; Wonser 9.3 21., 1938; Foster 93:. 3.1., 1939; see Appendix V) the recent natural *wallsc. send on Lake Nipissing surface: sass} Sec. 12 'r 39 N a 22', Rapid River Quadrangle, UBDI Geological Survey, 1958. (I [I 7S vegetation is reported to be mainly white and red pines. Personal observations by the writer confirm the existence of this vegetation type; however, two relatively undisturbed sites were found which were largely composed of hemlock, yellow birch and red maple.* There are no records, as far as the writer is aware, of pure hardw00d stands on Wallace semi. BROWN PGDZOLIC SANDS Only one soil series belonging to the Brown Podzolic great soil group has been adequately described in the yellowish sand soil area of inchigan. This series has been given the name Grayling arri is represented by only one soil type, Grayling sand. Recently, however, some sand soils origimlly identified as Grayling sand were found to have loany sand or sandy loam subsoil bands; these soils are currently being classified as Graycaln sand but little informtion is available on their distribution or range in natural vegetation. agree 2 Grayling sand differs from the Podzol sands by having an eluvial horizon less than 2 inches thick and by having an illuvial zone less than 2 feet thick. The illuvial horizons have yellowish (101R) colors or (bill (cln'ons less tlnn 6) reddish colors and overlie several feet of acid (pH values mually between 5 and 6) sand. This soil often grades laterally into Rubicon sand or into Croswell sand, a moderately wells-drained Podsol. *One of these sites is in the stuiy area (see previous footnote). The other is in Marquette County and was shown to the writer by Donald Buchamn (USDA Soil Conservation Service). 76 Grayling sand is usually found on ”dry sandy plains" or "the drier pine plains” (Mick, _e_t_ 91., 1951; Veatch, at 31., 1936). It occurs most frequently in glacial deposits (see Appendix V) and in Delta County it can be found in flmio-glacial deposits which were inundated by the later stages of Glacial Lake Algonquin. In Delta County, Grayling sand occurs only on surfaces which are over 3500 years old. The natural vegetation of Grayling sand in the Upper Peninsula consists largely of Jack pine, ”scrub oak”* and scattered red pine (see Appendix V). In the lower Peninsula, south of Indian River, white oak (guercus 93:13) and black oak (guercus velutina) are also forest components on Grayling sand. BRUNIZEM SANDS The only Brunizen soil series in the Podzol Region of Michigan has been given the new of Sparta and includes types which have all developed in well-drained sands. Sparta is quite restricted in its range in Michigan, occurring only in Podzol Zone I. It is described as having an A1 horizon 8-20 inches thick with a brighter colored illuvial (‘3) horizon immediately beneath (see Appendix V). Veatch (1938 and 1910) states that the Sparta series in Michigan developed in basim which were fornsrly occupied by shallow grassy or nrsh lakes. He postulates that these lakes dried up during a post- glacial "dry period.” *‘Scrub oak'I robebly represents rcus rubra v. borealis, ercus elli oidalis a or hybrids of sons e ems ea Quercus ra _ persoml cannunicat on ran . J. E. Cantlon, Botarq Department, Michigan state University). n 77 Hauser (1953) states that these areas are vegetated by assemblages of characteristic prairie species such as Andrgmgon gerardi, Andropogon ”Mus, Sorghgstrum m, Koeleria cristida, Eragrostis pectrmacea, Liatris m and Hieraciun longipilmn. However, the high incidence of are}; Efllnnica and certain weecw plants is not typical of other prairies. Many species in the Newaygo prairies are also found in the Sand Brrens of central Wisconsin but a few species are more typical of the Bracken Grasslands in northern Wisconsin, viz. §_o_a_ co ressa, £93 pratensis and M acetosella (1133 Curtis, 1959). The Bracken Grass- lands are in a great soil group transition zone similar to that in Nsseyge County but they occur on loans to fine sands and are not mapped as Brunizem. Curtis states, however, that the Bracken Grassland soils have a fairly deep incorporation of organic matter and little evidence of a highly leached A2 horizon. Part II. The Present Study 78 0mm 7. CLIMATE OF THE UPPER PENII‘EULA The cli-te of the Upper Peninsula was analyzed by means of water balance computation (according to the Thornthwaite System) and simmeries of man fall precipitation, mean annual snowfall and months of maximum precipitation. These variables were chosen because of their correlation with soil and vegetation characteristics in the Lower Peninsula. The data used in the analyses were obtained free U.3. Weather Bureau publi- cations (Weather Bureau, 1951;, 1958, and 195%). The detailed procedures used are the saw as those used to obtain the sale infor-ation for the Lower Peninsula (see Brunnschweiler, 1962 and Messenger, 1962) except that nonths of precipitation mum and co-nxila were determined on the basis that all other months had lean values at least 10% lower instead of 20% lower. The regloml distribution of precipitation regiaes for the Upper Peninsula can be seen in Figure 12. The southwestern section of the peninsula Ins a continental type of regim. The reminder of the peninsula has a regime which reflects a greater lacustrine influence on the climate with a September m or co-mxim of precipitation. The loan fall precipitation is plotted in Figure 1.3. The isohyets indicate that the western one-half of the peninsula receives less than 9 inches. Within that area, nest of Menominee, Dicldmon and Iron Counties receive less than 8 inches. Mean annual snowfall, shown in Figure 11;, is highest near lake Superior 3 from Ironwood through the szeemw Peninsula it averages over 120 inches. In southestern Delta County and throughout most of Menominee County, an annual snowfall is lees than 60 inches. Average annual PE values are plotted in Figure 15. The lowest values 79 80 . ZO_._.<._._Q_ommn_ EDEXSZ do zom> 4H_o< O as D O Q. . o v .mnEm>oz :_ : mm:em.o§...:om . u4m<.__<>< 3.325 2422 w_.oE. CHAPTER 8. DESCRIPTION OF THE STUDY AREA Seven well-drained, relatively undisturbed sandy sites in Delta County and one in Alger County, Michigan (see Figure 19A) were chosen for a study of: (1) their soil characteristics and (2) their inter- relationships to each other and the soil-formation factors . ans-n (The clinte of the study area is strongly influenced by the posi- tions of Lake Michigan and Tales Superior. Segments of the area which are frequently exposed to lake-altered air masses exhibit this effect most clearly. During the months from May through October the prevailing wind is fro. the south at Escanaba. This lake breeze has a moderating influence on the temperature of Escanaba and probably much of southern Delta County. During the months from November through April the prevailing wind at Eeoamba is from the north or northwest. Thus the winter temperatures in southern Delta County are less moderated by the lakes than are the 8Il-er telperatures. Green Bay, Little Bay de Noe and Big Bay ds Noe r:Iseese over during thewinter, and during the part of thewinterthey fire frozen, the lake influence on temperatures is even less pronounced. The open water of lab Michigan proper, however, appreciabh moderates the winter te-pereturee in the Fayette-Sac Bay area in the southern Pill-t of the Garden peninsula at the southeastern tip of Delta County. For the period 1931-52 the mean temperature, precipitation and “fiber balance values for Chathaa, Escambe and Fayette-Sac Bay are Presented in Table 5. 88 89 Pmdmo uJOOIUm 03Pm 20m. 90 TABLE 5. SOME CLIMATE FEATURES OF THE DELTA-ALGER STUDY AREA (1931-52)* Escenabe Chathss Fayette-SB Escalate Chatlna Fayette Escansbs Fayette Escambe Chathsa Fayette WURE (°F) J F n 1 n J J 1 s 0 N DAVE 18.6 18.3 26.3 38.6 50.1: 61.0 67.3 65.3 57.5 h7.3 311.0 25.9 112.5 17.6 17.1. 21»? 37.8 h9.8 60.1 65.9 617.1 56.6 h6.1 32.2 21.9 1.1.2 20.3 19.6 27.0 38.5 h9.3 59.0 66.1 65.3 58.1; h8.0 35.h 25.2 h2.7 mscrpmxon (13cm) J F n 1 M J J 1 s 0 N DTOTAL 1.61 1.29 1.68 1.95 2.86 2.96 3.514 3.03 3.01; 2.09 2.30 1.37 27.72 2.30 1.68 1.72 2.08 3.00 3.68 3.27 3.37 h.22 2.81 3.39 2.20 33.72 2.10 1.68 2.12 2.31 3.05 3.113 3.31; 3.00 3.39 2.1:6 3.20 1.99 32.07 snonm (mans) J F n 1 n J J 1 s o N DTOTAL 1h.6 12.7 9.7 2.8 0.1 0.0 0.0 0.0 0.0 0.2 h.9 10.1: 55.1: 19.1: 13.6 10.8 6.1; 0.6 1' r T 'r 2.6 12.7 17.9 8h.0 20.7 18.8 114.3 5.6 0.1; 0.0 1' 0.0 1' 0.5 8.7 16.1 85.1 mm sums (WEBB 03' um) IN son. Inn 111- CAPACITY IN son. wrrH h' wmm P3 DEF mm suams NOVEMBER SURPLUS (candy 10!!) (and: loan) (and) 558 15 161 o 553 10 33h 90 51:6 11 280 to *rhe raw data for this table were derived from U.S. Weather Bureau publications (weather Bureau, 1959a and h). These data show that Escamba has clintic aodsture characteristics noretypicelofPodzol ZoneIIthanPodzolhnem. Basedonthese 91 data and Figures 13 and 124, it is apparent that Site Bl south of Escanaba, Figure 19, exists under a somewhat less favorable climate for dark upper illuvinl horizon develoment than do the other sites in this study. Surface Geolog The surface geology of Delta County and neighboring counties is depicted in Figure 1913. Delta and Alger counties were completely covered by Valders ice (Bough, 1958). The unadulterated outwash plains occurring in these two counties, then, should be of Valders age. However, since these counties lie about 200 miles north of the point of mxinum advance, Valders drift is logically somewhat younger in these northern areas tlnn it is farther south. is a nutter of fact, Hough postulates that the ice was still present in Delta County during the highest stage of Glacial Lake Algonquin. Accordingly, then, the longest period of tine that any surfaces in Alger County and northeastern Delta County have been exposed to post-glacial neathering should be about 10,500 years (Broecker and Fan-and, 1963). lake Algonquin was assumd by Leverett and Taylor (1915) to have extended into the Lake Superior basin through the Au Train-Whitefish Valley system in central Alger and Delta counties. Observations of lacustrine features were nude by those workers, and subsequently by others, up to 960 feet above sea level in the vicinity of Munising in Alger County. The highest of these features was much higher than the isobase line of the highest Algonquin level derived by leverett and Taylor fron observations in other localities. As a result, these features were assumed to be of local origin. Bergquist, working in this region in the 1930's, also assumed these features to be local in FIG. l9-B SURFACE GEOLOGY OF DELTA AND ALGER COUNTEfi SENEY COMPLEX INDIAN LITTLE BAY DE LEGEND Moramol Deposns 0 000 ’ 7 - ‘0‘“ TI“ Plum Deposns @ Sandy Lake—Plain Deposits A Muck and Peat Over Lucustrine Deposits [j Drumlin Deposits Dune Sand v Miles io‘fl} Outwosh Bedrock O 2 4 ‘ I_1__A_A_J L-S_°_,‘fl'3___s_‘_"_f_'9l"960 8 Bergquus',l936 Z6 93 origin since they did not correspond in elevation to the highest lacustrine features farther to the east. South of Munising in southern Alger County, and northern Delta County, observations of lacustrine features are lacking. Based on this dearth of observations, the anonlous lacustrine features near Munising and the configuration of the moraine-outnsh system in that area, Hough postulated that: (l) the highest stage of lake Algonquin did not extend into the lake Superior basin because it was damned by glacial ice and (2) by the tine the Au Train—Whitefish valley system we ice-free, Algonquin labs waters were at such a low level as to prevent a lake connection from occurring between the lake Superior and Lake Michigan basins in that valley. , Glacial lake Duluth, which is assuned to have been contemporaneous with the min Algonquin stage but confined to the western part of the lake Superior basin, is thought by Hough to rave used the Au Train- Ihitefish valley system as its spillway following the unblocldng of this passageway by the glacial ice during the "Upper Group” of lake stages in the Lake Michigan basin. The resultant deluge is postulated to have inundated all but the highest moraines in Delta and Alger counties. During further retreat of the ice front to the north and east, the water levels in both of the above-mentioned basins dropped to a very low elevation, tm lowest of these in the Ialne Michigan basin being Lake Chippewa. This subsidence occurred between about 10,500 and 9570 2 1.50 years ago (Breecher and Farrand, 1963). 1'01le the lake Chippewa period, water levels rose, and presunbly by 5720 2: 250 years B.P., they were close to the present- I. 98 day levels (based on the buried ash tree referred to in Chapter 3). By 3500 years B.P., a stable level was reached in both basins. This level has been given the name lake Nipissing and the present elevations of its beaches rise from 609 feet above sea level at Escanaba to 629 feet at Munising (Leverett and Taylor, 1915). Following lake Nipissing tines, water levels dropped about 10 feet and became stable enough at that point to be classified as a separate lake stage. The name given to this stage was lake Algona and it has been dated at about 2500 years B.P. (Hough, 1958). lake Algoma beaches and terraces have been identified on Garden Peninsula (eastern Delta County) at elevations of 590 to 600 feet (Leverett and Taylor, 1915; Bergquist, 1936). In Schoolcraft County, further east, beaches are also present which are transitional between the Nipissing and Algoma levels. Bergquist states that these beaches were formed either during the lowering of the Nipissing waters or by store waves of lake Algosa. The Algosn shore in Schoolcraft County is not traceable as a continuous feature but occurs rather as more or less disconnected units along the lake Michigan shore. In certain areas, the Algoua features are very definite but in the main they are either obscured by low data developments or are nissing entirely as a consequence of later wave activity. In several places a series of low fore-dune ridges or limestone rubble ridges extend outward from the Algona shore features to the present shore. Bergquist states that these ridges my represent store wave deposits of the present lake. Stern waves are maples of short-period fluctuatiom in lake levels. The main rise resulting frat short-period fluctuations that has been observed at gage sites on lake Michigan is 2.8 feet at 95 Calumet Harbor, Illinois. However, the ability of the wind to raise water levels is greatest in bays and extremities of the lake especially when the wind is blowing toward these locations. At the east end of lake Erie, for imtance, the maxim short-period rise recorded was 8.1; feet (laidly, 1962). The monthly average level of lake Michigan reached the maximum recorded value of 583.6 feet above sea level in June, 1886. This peak mked the end of a four-year period during which the monthly average level exceeded 583 feet each summer. No other comparable period has been recorded (laidly, 1962). The writer has identified what he believes are Algosa terraces cut into linestone bedrock at several locations on Stonington Peninsula and on the east side of 0gonts Bay in Delta County. These terraces invariably occur between the 590 and 600-foot contour lines. In some areas, terraces or low, dune-like ridges are present above Algona Inve- cut cliffs and below lake Nipissing terraces. In several areas, a series of parallel, low, fore-dune ridges extend outward from the base of Algoma wave-cut cliffs to the present beach. A very conspicuous area of ridges such as these is to be found at the head of Big Bay de Noc although no Algon cliff was identified at the upper end of that series of ridges. Sit: Locations Three sites are located on low dunes below lake Algonn wave-cut cliffs. These dunes are considered to be post-Algom in age based on their similarity to those described by Bergquist and since buried soils are present beneath a relatively unleached sand cap varying in thickness from six inches to a foot. Excavations in the vicinity of one of these sites (Site Al) revealed nan-made wood chips and a buried tree trunk 96 beneath the sand cap. Ages of two of the larger trees growing on each of these three sites did not exceed 70 years. Since lake level records for Lake Michigan indicate that from 1882 through 1886 the monthly average lake level during the peak months (summer) was between 583 and. 581; feet, a portion of the surface now beneath the sand cap would have been inundated and storm waves say have inundated most of it. Since loggixg began in the general area between 1880 and 1890, the sand cap on these surfaces was probably deposited no earlier ttnn 1880 and in all likelihood was deposited during and immediately following the high lake levels in the add-1880's. Until further investigations are ands, these sites will be collectively referred to as being post-aAlgom in age. These three sites are designated as Al, A2 and A3 with the numbers varying directly with the apparent degree of Podzol development. Site Al is about 100 yards from the present beach and is adjacent to the dune heath (ninly JuniErus spp.) sons. Site AZ is about 200 yards inland and is within a few hundred yards of Site Al. Site A3 is about 16 miles northeast of Sites Al and A2; it is also about 200 yards inland. Sites Al and A2 are bounded on the south and west by water and on the north by a boggy area. Site A3 is bounded by water to the south, by beg to the east and northeast, but by well-drained uplands to the west and northwest (site locations are shown in Figuresl9A and B). Tm sites are located at levels between those of the A sites and lake Nipissing terraces. The parent naterial at each site is composed of a stratum of medium to fine sand overlying a calcareous stratum of coarse sand, gravel and gastropod shells. These sites are designated as Bl and B2 on the basis of a darker upper illuvial horizon in the B2 9? soil. The Bl site is bounded by moist sites and is about 100 yards from the Bark River. The B2 site is bounded by dry sites and is about 500 yards northwest of Site A3. Two sites are located in the area which was inundated by the waters fron Lake Duluth as they spilled southward into the Lake Michigan basin. Both of these sites lie well above the highest Lake Nipissing features therefore the land surfaces are of Sub-Duluth (Hough, 1958) age. These two sites are designated as 01 and 02 on the basis of a darker upper illuvial horizon in the 02 soil. Site 01 is bounded on the west by an extensive lowland area which begins at the bottom of the escarpment which leads down from the terrace on which Site 01 is located. Site 01 1: about 100 yards fron the escarpment. Site c2 is surrounded by well-drained sands with occasional pits typical of outwash plains. Both of these sites lie inland more than 10 miles fmn either Lake Michigan or Lake Superior. Site D1 is located near the top of a moraine which apparently was not immdated by the Sub-Duluth deluge. This moraine is a part of the Newberry Morainic System (Bergquist, 1936), and as such, would be of Valders age. Site D1 is well inland from the Great Lakes but is situated about 3/h of a mile south of an extensive area of smll lakes, all of which lie at elevations 100 to 130 feet lower than Site D1. The approximate ages in years of the surfaces at these sites are therefore assumed to be as follows: 9? soil. The B1 site is bounded by moist sites and is about 100 yards from the Bark River. The B2 site is bounded by dry sites and is about 500 yards northwest of Site A3. “0 sites are located in the area which was inundated by the waters from lake Duluth as they spilled southward into the lake Michigan basin. Both of these sites lie well above the highest lake Nipissing features therefore the land surfaces are of Sub-Duluth (Hough, 1958) age. These two sites are designated as Cl and C2 on the basis of a darker upper illuvial horizon in the C2 soil. Site Cl is bounded on the west by an extensive lowland area which begins at the bottom of the escarpment which leads down from the terrace on which Site 01 is located. Site Cl is about 100 yards from the escarpment. Site C2 is surrounded by well-drained sands with occasional pits typical of outwash plains. Both of these sites lie inland more tkan 10 miles from either lake mchigan or lake Superior. Site D1 is located near the top of a moraine which apparently was not inundated by the Sub-Duluth deluge. This moraine is a part of the Newberry Morainic System (Bergquist, 1936), and as such, would be of Valders age. Site D1 is well inland from the Great Lakes but is situated about 3/h of a mile south of an extensive area of smll lakes, all of which lie at elevations 100 to 130 feet lower than Site D1. The approximate ages in years of the surfaces at these sites are therefore assumed to be as follows: SITE A2 B1 32 Cl 02 D1 98 AGE-ROUGH CHRONOLCBY post-Algom (writer's estimate) post-Algona (writer's estimate) post-Algoma (writer's estimate) 3000 (writer's estimate) 3000 (writer's estimate) 7500 7500 8500 AGE-BROECKER AND FARMND CHRONOIDGY 10,000 10,000 10,500 CHAPTER 9. METHODS USED IN ECOSYSTEM INVESTIGATIONS m Couposition Forest composition at each site was determined by the use of a basal area prism with a soil pit as plot center. In these descriptions, the following terms are used: (1) ”dominants" which refers to those trees having their crowns above, or at the same level as, crown of the neighboring trees; (2) "intermediates" which refers to those components from one inch in diameter (at breast height) up to, but not including, the dollmnt trees 3 (3) “reproduction” which refers to tree reproduction and includes seedlings and saplings up to one imh in diameter. In the individual site descriptions, these term are abbreviated to D, I and R. aw... Ens: The humus types of the profiles studied are classified according to the system developed by the Comittee on Forest Humus Classification, Forest Soils Subdivision, Soil Science Society of Alerica (Hoover and Lunt, 1952). These designations are cited preceding each profile description. The intial appearance of the 2 33.33 Rubicon sand hum layer, Site 01, suggested a non humus type. Closer inspection, however, revealed that what appeared to be an H-lsyer was actually a fixture of bleached sand grains and black humus which graded rather abruptly into the underlying Ea (A2) horizon. The estimation of the percent organic ntter based on organic carbon content revealed that this dark horizon contained only about 7% organic setter therefore the humus layer seemed to be most appropriately classified as a Shallow Sand Hull. There was some question as to whether the upper hundc horizon of Site A3 met the specifications for an H-layer since it contained 99 ,‘l 100 scattered, bleached sand grains. Since this horizon was found to contain 29% organic matter and graded gradually into the Vh horizons below, the upper hunts horizon was considered to be an H-layer and the humus type was classified as a thick duff-mull. In a recent paper, White (1965) states that there is no evidence of biological incorporation of organic setter in some lake States humus types which morphologically appear to be duff-mulls. The V'h (A1) horizons underlying the Oh horizons of such soils usmlly contain less than 14% organic netter and therefore it was felt that such humus types should be classified as more rather than duff-nulls. It is possible that the humus layer of Site A3 is also of this type but confirming data other than z 0.1!. is lacking. Comequently, the humus layer is tentatively classified strictly according to its morpholog. The humus layers of Profiles A1, B2 and D1 definitely lack an H- layer and exhibit a gradual decrease in organic matter with depth. 111 Profile D1, however, there was a gradual decrease from the Vhl to the Vh2 and then an abrupt decrease from the Vh2 to the Vh3. It seems plausible that the Vh3 contains mostly infiltrated rather than biologi- cally incorporated organic setter. In Profile B2 also, the lowermost Vh horizon say contain mostly infiltrated organic matter. Since these problems cannot be resolved with the data at hand, these humus layers are classified as mulls. Due to the looseness and low organic matter content of the humus layer in Profile Al, it is classified as a sand mull. The other two are classified as coarse or median nulls. The lack of an on horizon on a well-developed Podzol may be a residual effect from disturbances during earlier lumbering Operations . 101 However, the height of the trees and the density of the stand are such that the writer feels that an 0h horizon would have subsequently developed if the foliar composition of the stand were conducive to such a development. Soil Profile Descriptions Soil profile descriptions were made at the time of sampling (July and August, 1961). Horizon designations are those suggested by Whiteside (1959). Designations for the horizons in the following profile de- scriptions have the following equivalents in the nomenclature outlined in the 1962 supplement to the Soil Survey Mamnl (Soil Survey Staff, 1951): Of = 01 (F layer) 011 = 02 (H layer) Vh -"-' Al (and H layers containing less tmn 20% organic natter)* El 3 L2 1111b 3 Bhir Bibi = Bhir lhbic = Bhir (or ma if cemented horizon is at least 90% continuous) lib = Bir 1131 I Bir w = C (or Cl formerly) P = c (or 02 formerly) U = II C (or formerly, D) p = b (buried soil) “The above mentioned supplement stipulates that organic horizons of mineral soils should contain more than 20% organic matter if the mineral fraction has no clay. This criterion was used to separate on from Vh horizons in this study. 102 colors are for moist soils; color names are those of the ISCC-NBS (U.S. National Bmau of Standards, 1955). Acidity was determined with a Hellige-Truog pH kit (pH meter values are listed with the soil test data). SITE L1: SAND REGOSOL UNDER BALSAM FIR Location: sass}, Sec. ll, Twp. 38N, Rge. 22w Dreimge: well-drained (water table at 7 feet in summer of 1961) Slope an! aspect : negligible Topography: slightly undulating to level Landforms low dunes . Elevation: between 580 and 590 foot contours on topographic nap Forest characteristics : basal area/acre = 90 sq. ft.3 balsam fir - 89%, DR 3 paper birch - 1.1%, D. Hume type: deep sand mull Field description of horizons : Horizon Depth (mg Characteristics 0: +1 to o balsam fir debris; white fungal mom; pH variable, spots of 5.0, 6.5 and 8.0. Vhl 0 - 1 sand; black; pH 8.0. M l - 14% sand; brownish gray (1013 3A)! PH 8.0. p h% - 10 sand; light yellowish brown (101R 6/14); pH 8.0. pVh 10 - 114% sand; dark grayish yellowish brown pIib lite - 175 sand; moderate yellowish brown (lam 5/103 pH 8-0- P 17% - 66 sand; light yellowish brown (1013 6/10; pH 8.0. 103 SITE A2: EASTPORT SAND UNDER PINE Location: swede}, Sec. 12, 'I‘Ip. 38N, Rge. 22w Drainage: well-drained (water table at 5% feet in summer of 1961) Slope and aspect: negligible Topograplv: slightly undulating to level Iandfora: low dunes Elevation: 590 feet Forest characteristics: basal area/acre = 1140 sq. ft. 3 white pine - 29%, D; balsam fir - 21%, I; aspen - 21%, D3 white-cedar - 21%, I; red pine - 7%, D; paper birch - 7%, D. Hunts type: tl'dck nor Field description of horizons : Horizon Depth (1m) Characteristics or +23,- to +1% mostly coniferous debris ; yellow and white fungal melts; pH variable, spots of 5.0 and 6eSe 0h +13; to 0 black humus with scattered, bleached sand grains 3 pH h.0. E, o .. 1% sand; grayish yellowish brown (10m 5/2); pH heo "’ Seoe lib ll} - 6} send; strong yellowish brown (101m 5/6); pH 5e2e ‘ I/pV 6% - 12} send; moderate yellowish brown (10m h/B); pH 3.0. P 12‘} - 66 sand; light yellowish brown (101R 6/10; pH Boo. 10h SITE L3: EASTPGRT SAND UNDER RED OAK Location: NWiSEfi, Sec. 2h, Twp. hON, Rge. 20W Draimge: well-drained Slope and aspect: negligible Topograplw: sligltly undulating to level Landforl: low dunes Elevation: between 580 and 590 foot contours Forest characteristics: basal area/acre = 120 sq. ft. 3 red oak - 67%, D3 white pine - 25%, 1); sugar maple - 8%, DIR. Humus type : thick duff-mull Field description of horizon: : Horizon Depth (mg Characteristics at +13; to a pine and hardwood debris ; white fungal melia; pH 6.2. 011 0 - 1 dark grayish yellowish brown to brownish gray (lOYR 2/1-3/1) humus; scattered, bleached sand grains; pH 5.0. Vhl 1 - 1‘} brownish gray (101m 3/1) humus and sand mixture; PH heOe Vh2 1%; - 2i sand; light brownish grey (1013 5/1); pH 13.0. as 2} - hi sand; light grayish brown (7.513 6/2); PH Seoe l'ib his - 101; sang; strong yellowish brown (101m 5/6); PH eze pVh 10} - 11-3/1: sand; brownish grey (101R h/l); pH 6.0. pm 11-3/1; - sand; light grayish yenowish brown 12-3/h (1013 6/3); pH 6.0. pIib l2-3/h - sand; strong yellowish brown (101R 5/6) 3 16.3”! PH 6e0e I 16-3/14 - 38 sand; light yellowish brown (101R 6/14); pH 7.0. P 38 - 66+ sand; light yellowish brown (101R 6/h); pH 8.0. 105 SITE Bl: RUBICON SAND UMBER HELEOCK Location: NEfi‘l‘IEi‘, Sec. 27, Twp. 37N, Rge. 21m Drainage: well-drained Slope and aspect: zero Topography: slightly undulating to level landforn: low dunes ("Upper Algoma”) Elevation: 595 feet Forest characteristics: basal area/acre = 220 sq. ft. 3 eastern hemlock - 36%, DIR; yellow birch - 23%, D3 white pine - 18%, D; paper birch - 11%, D; white spruce - 5%, D3 balsam fir - 1%, DR; red maple - O, R. Humus type: thick mor Field description of horizom : Horizon Depth (ind Characteristics 01' +3 to +2 Gymnosperm and Angiosperm debris; white fungal melia; pH variable, from h to 7. 0h +2 to 0 black to dark gray humus with scattered bleached sand grains 3 white and yellow fungal mycelia; pH 11.0. Vh o - é send; grayish brown (5m h/2); pH h.o. an i - 7% and: light mesh brown (7.5312 6/3); pH the. 11:11: 7% - 98‘ sand; strong brown (511?. 14/8); pH h.0e Iib 9% - to sand; strong yellowish brown (7.513 5/8); pH 6.0. P to - 66 sand; light yellowish brown (lOYR 6/h); pH 8.0. U 66+ sand, gravel, gastropod shells and shell fragments ; pH 8.0. 106 SITE B2: EAST IAKESANDUIDERRED OAK Location: SEfiNWfi, Sec. 21:, TWp. hON, Rge. 20W Drainage : well-drained Slope and aspect: 3% east Toposraphw: low dunes ("Upper Algom") 595 feet Iandform: Elevation : undulating Forest characteristics: basal area/acre -"- 130 sq. ft. 3 Humus type: red oak - 5h%, DI; sugar maple - 31%, DIR; beech - 8%, DI; basswood - 8%, DI. Field description of horizons : Horizon Of Vhl lib d Depthg(in.) 4-1/8 to 0 0-2 2-5% 52-6 but mostly absent Sit-8’: Bat-15% si-a 26+ very deep coarse or medium mull Characteristics hardwood debris; pH 7.0. black hunms and sand nurture; pH variable, spots of h.0 and 6.5. send; brownish gray (101m 3/1): pH 11.0. sand; grayish yellowish brown (101m 5/2); PH heSe send; strong brown (513 3/6): pH h.8. sand; strong yellowish brown (7.5m 5/3); pH 6e00 sand; dark orange yellow (101R 6/6); pH 6.0. coarse sand, gravel and gastropod shells; light yellowish brown (101R 6/h); pH 8.0. 107 SITE Cl: RUBICON SAND UNDER RED PINE Location: Minna», Sec. 7, Twp. h2N, age. 20w Dreimge : well—drained (water table deeper than 11: feet) Slope and aspect: negligible Topograplv: level Iandform: plain Elevation: 750 feet Forest characteristics: basal area/acre -"-'- 160 sq. ft.; red pine -- 100%, D 3 red maple - 0, R; balsam fir - O, R. Humus type: shllow sand mull Field description of horizons : Horizon Depth (in) Characteristics 0f 2% to 0 pins debris 3 white fungal nycelia; pH 11.5. W: o - 1} black humus and sand manta-e; yellow fungal melia; pH h.0. ms 1% - 6 send: light grayish brown (513 5/2); pH 11.0. lhib 6 - 13 end; moderate brown (7.5m h/h): pH 5.0. Iib 13 - 35 sand; strong walla-i111 bro-n (7.5m 5/6)s pH 6.0. n 35 - 156 send; light brown (7.5m 5/h); pH 6.0. 5 156 4- fine sand; light brown (51H 5/3); pH 6.5. 108 SHE CZ: KALKASKA SAND UNDER HEMLCBK Location: xvi-m, Sec. 10, Twp. hsN, Rge. 19w, Alger County Michigan Draimge; sell-dram (water table deeper than 1).; feet) Slope and aspect: 3% south Topography: undulating Landforms outwash plain- Elmtion: 900 feet Forest characteristics: basal area/acre = 160 sq. ft.; hemlock - 58%, DI; yellow birch - 21%, D 3 rod mph - 110%. 1: balsam fir - 7%, I. Humus type: thick mor Field description of horizons : Horizon Depth Lin .) Characteristics Of +3 to +2 coniferous and hardwood debris; white fungal mom; pH variable, 14.0 with spots of 6.0. on .2 to 0 black humus; yellow fungal mocha; pH h.o. h 0 - 6 sand; light grayish brown (7.5m 6/2); In 11000 Ihib 6 - 8 sand; dark: brown (SIR 2/3); pH 14.0. nlbic 8 - M; sand; discontinuous, indurated tongues; dark grayish brawn (SIR 2/2); pH 5.5. 1131 8 - 26 sand; strong yellowish brown (7.513 5/8); pH 5.50 If! 26 - 38 san§35rtrong yellowish brown (7.515! 5/6); pH 0 e I 38 - 96+ sand; light brain (7.5m 6/h); pH 6.0. 109 SEE D1: KALKASKA SAND UNDER SUGAR MAPLE Location: MM, Sec. 7, Tip. hBN, Rge. 18W Draimge; well-drained (water table deeper than 11; feet) Slope an! aspect; Id east Topography: rolling Iendfm: moraine Elevation; 890 feet Forest clarecteristics: basal area/acre = 180 sq. ft.; sugar mp1s - 89%, DIR; black cherry - 11%, DR. Hunus type; very deep coarse or medium null Field description of horizom : m Depth (2111.) Characteristics m o - it black humus and sand mixture; pH 5.0. m t - l sand; very dark gray (513 3/1); pH 6.0. We 1 - 5 Band; dark gray (SIR M1): pH 5.0. h 5 - 7% and; rOddiBh my (SIR 5/2)3 pH 5.8. Ihibl 7% - lO sand; dark reddish brown (513 2/2); pH 6.2. Ihihz lo - in sand; reddish brown (513 MB): pH 6.8. Ibl 1h - 31% Bands strong bra-n (7.5m 5/6): pH 5.5. ll 31; - lhh sand; yellowish bro-n (7.5m S/h); pH 6.0. W2 1104-168 sandwith thin, orange to pink loony and bands; sand—brown (7.5m 5AA); acid. 223.122 Eng Three Lakes Bog is located approximtely 100 yards south of Site 02. The bog is surrounded by sand for at least 3% miles in every direction. The soil type surrounding the bog for at least 3; mile in every direction 110 is Kalkaska sand (Veatch, 2+: 114., 19314 and confirmation by writer). The nrgin of the bog is bounded by steep slopes; consequently well- drained soils are encountered within a few feet of the bog margin. The bog contains 9% feet of peat overlying sand into which some organic mtter has been incorporated. The surface of the bog is completely covered with a mt of sphagnum interspersed with leatherleaf. The forest composition immediately around the bog contains a somewhat higher percentage of red maple and alder (A_l_._n_u_s ma) than is found farther from the bog margin. Tree species found in the upland areas around the bog which are not included in the Site 62 description are white pine, American beech and white spruce. Sugar mp1s was notably absent in this forest cover type which was designated as hemlock on the 1951i U.S. Forest Service Tinber Survey map of Hiawatha National Forest. Sampling of the peat bog was accomplished with the use of a Hiller borer by Dr. A. T. Cross of the Geology and Botany departmnts, Michigan State University, and the author. Samples were collected from three- to six-inch intervals within the peat itself and as a continuous core for the first six inches of the underlying peat—sand mixture. The samples were kept frozen until macerations were begun and kapt under refrigeration between subsequent sub-samplings . The first maceration technique employed was simply the boiling of the wet sub-sample in 10% NHhOH. This treatment did not isolate the pollen grains sufficiently, therefore bleaching with sodium hypochlorite prior to the 1!th treatment was tried. This treatment, even in dilute concentrations, destroyed all the birch pollen. Sodium chlorate, how- ever, proved to be safe to use (standard treatment used by several palynologists; see Brown, 1960). 111 The technique finally employed was as follows : (l) About an inch of wet peat was loosely packed into a ho ml centrifuge tube and 10% mm was added until the peat was completely covered. The tube was then placed in a 100°C water hath for % hour with occasional stirring. Next the fixture was centrifuged at 2500 rpm for 5 minutes and the supernatant liquid decanted. The residue was then washed with water and centrifuged with subsequent decantetion. The residue was next dehydrated with glacial acetic acid followed by centrifugation md decentetion. (2) For lignin oxidation, 5 ml of glacial acetic acid, 5 I]. of 50% Na0103 and 1 ml of concentrated HCl was added to the centrifuge tube and the mixture stirred and allowed to bleach. The tube was then filled with glacial acetic acid, centrifuged and decanted, then washed twice with glacial acetic acid with centrifugation and decantetion. This technique is recommended by Faegri and Iversen (see Brown; 1960). (3) For cellulose decomposition, an Erdtnn (see Brown, 1960) acetolyeis mixture (9 parts acetic anhydride to one part concen- trated 3250‘) we added to the residue, and the mixture placed in an 80°C water bath. The bath was then brought to the boiling point and allowed to boil. for two minutes. The tube was then centrifuged, the liquid decanted and the residue washed with glacial acetic acid followed by centrifugation and decantation. The residue was finally washed with water (followed by centrifugation and decantation) until a clear eupermtant liquid was obtained. The resultant residue was then strained through a double layer of cheese cloth with concurrent washing. The suspension was concentrated by centrifugation and transferred to vials frm which slide preparations were made. Pollen counts were made on glycerine mounts only. Size frequency 112 distributions were nude on pine pollen grains using the distance between upper wing imertions as the criterion. The resulting bar graph indicated a ml curve distribution around a distinct hit-micron peak and a suggestion of another peak at SS nicrom. The hit-nicron peak was at- tributed to the presence of Jack pine and red pine pollen based on the stub of Cain and Cain (191:8). Based on the presence of a "saddle” in the distribution curve at 50 microns, .11 pine pollen grains having dinemiom greater than 50 microns were considered to be from white pine. Two hundred tree pollen grains were counted per slide. Non-arboreal pollen was not considered except to note that it was not prevalent at any level. Folder images A cuposite foliage sample was collected from one tree of one or acre representative species on each of five plots. The samples were collected between 9 a... and 12 noon during the first two weeks of SepMer, 1961. Collections were made from the southern, lower one- third of the crown only. In the case of conifer foliage, only the current year's growth was sampled. These-plesweresubjectedtooven-drying onthe myoftheir collection; drying was accomplished at 70°C. The oven-dried ell-plea weregroundinalileynfllandthenturnedovertobni. L. Kenworthy of tin Horticulture Departmnt at Michigan State University. With the emception of total nitrogen, the elemental analyses were done spectro- DOOM e Soil Microflera Studies The hum-containing horizons free all A, B and 0 sites were ealpled to obtain a quantitative estinte of the bacteria, actinomcetes 113 and fungi. he samples were taken from each horizon; one sample was frozen for later determination of moisture content and nitrifying capacity; sub-samples were taken from the other sample for slide preparation. Slides were prepared within 21; hours after the collection of the samples 3 the preparation technique used was that of Jones and Mollison (l9h8). Three slides were prepared from each sub-sample; counts were made by the use of random traverses. Stained bacterial cells were counted with no regard for cell-size differences. In the case of fungal mphae, however, four diameter classes were established and length measurements were made with the use of a calibrated ocular grid; separate counts were made for unstained hyphal fragments. ictinowcetes were enumerated by length measurements of stained fila- ments only. Total Soil Carbon and m Matter Determinatiom Total carbon was determined in duplicate by the wet combustion method of Allison (1960). The values obtained can be interpreted as organic carbon except for those of the Vh2 horizon of Profile Al; this horizon contained carbonates as well as organic matter. 3112.5 Density 9; §9_i_l_ Horizom In order to evalizate the total amounts of organic matter (and other constituents) as well as concentrations it was necessary to estimte the bulk density of the various horizons. This was done by weighing the quantity of dry soil necessary to fill a sampling spoon calibrated to hold 2.5 gram of a soil raving a bulk density of 82.6 lb./cu. ft. or 2,000,000 lb./acre furrow slice (AFS). The weight of soil per AFS was calculated by the ratio: 111; ngt. of samle ' 2.5 TDJAFS 3 T—‘6,000,00 The percentage of the various constituents times the weight of soil per AFS (in pounds) then gives the pounds of the constituents per AFS. Pounds of constituents per horizon-acre (#/HA) :- thickness of horizon z I/AF'S. The thickness used for P, W and U horizons was determined by subtracting the solum thickness from 66 or 96 inches. Although sampling at 96 inches was only done in the case of Profile 01, visual examination and pH readings in the field did not indicate the necessity for sampling it this depth. 3939: _s_oi_1_ Nitrogen Determimtions Total nitrogen was determined in duplicate by the KJeldahl method essentially as described by Jackson (1958), except that no selenium was used. §£_i_l; m m Fractionation Allan-soluble organic nutter was extracted and fractionated accor- ding to the method outlined by Stevenson (1960). The lignin of the buds acid fraction was subjected to a theme-decomposition process (see Johnston, 196b,) and the products of decomposition were determined by paper and gas chrontomwu". No analyses were made on the fulvic acid fraction. Creme Acid g Nitrate Production of Soil Horizons The (21, VII and ll: samples used for this study were frozen shortly *Degradation and chrmtograplv was performed by Dr. Harry H. Johnston of Wilmington College, Iilnington, Ohio. 115 after their collection and kept frozen until sub-samples were taken. In the laboratory, either 5 or 10 gram sub-samples were placed in carbon filter tubes and leached to determine the initial nitrate content. Leaching was accomplished by using 60 ml of distilled water and vacuum, then adJmting the leaclmte volume to 60 :11. One to four ml aliquots were taken from the leachate for nitrate determination using phenoldi- sulfonic acid (Stanford and Hanway, 1955). The leached sub-samples were brought to a constant moisture tension by using full vacuum; then they were transferred to an incubator at 30°C. Following the initial incubation period, the sub-samba were leached at weekly intervals with 60 ml of distilled water and then re-incubated. The water-soluble organic letter (designated as crenic acid) appearing in the leachstes was measured relatively by determining the percent transmittance of the leachate at a wave length of 370 my. (medium absorption was actually in the ultra-violet, beyond the range of the calorimeter used). A Beckmn glass electrode pH meter was used for determining the acidity of the leachates. Soil treatments consisted of: (l) inoculation with a l:l0,000 suspemion of an actively nitrifying garden soil 3 (2) fertilization with 50 p}: P (as ordinary super phosphate) and 0.25% Ga (as lime); and (3) both of the above treatments combined. Two replicates of 10 grams each were used for the controls and single 5-gram samples were used for the treatments. Iron and aluminum were extracted from duplicate lO-gram sanoles of soil by the sodium dithionite—citrete-bicarbonate method (Jacks on and Mehra, 1960). An aliquot of the extract was taken and the organic matter 116 therein digested in a ternary mixture of concentrated acids (100 ml HNOB, 10 :1 stoh and ho ml HGth). Heating was continued until the solutions were evaporated to dryness. The resulting white residue was taken up in dilute (1!) HCl and brought up to 50 ml with distilled water. Aliquots were then taken for iron and aluminum determinations. Iron was determined by the KSCN colcrhatric method (Jackson, 1956). um was determined by the 8111an method using the modifi- cations recomended by Franzmeier (1962) in his work with similar soils. Available Phosphoms" _i_._n _S_9_i;|._ Horizons Available phosphorus was extracted from a 2.5 g sample of soil (approximately 2.5 g in the case of horizons below the upper humus horizon) with 20 ml of 0.03g nabs mind with 0.0253 H01 (Bray and Kurtz, 19115). The suspensions were skahen for one finite and then filtered. Phosphorus in solution was determined colaimtrically using the ammonium nolybdate—lwdrochloric acid solution of Dickson and Bray (l9h0) and the l-amlne, 2-mphthol, h-sulfonic acid reducing agent developed by Fiske and Subbarrew (1925). Exchangeable 1252* in _s_eg Horizons Mhangeable bases were extracted by adding 20 .1 of neutral 0.13 Mama to 2.5 g of soil (approaimtely 2.5 g in the case of horizons below the upper humus horizon), sinking the suspension for one minute and filtering. Calcium, mgnesium and potassium were determined on the extracts using a flame photometer. *Deterninations nude by Soil Testing laboratory, Soil Science Department, Michigan State University. 117 Reaction* _<_>_f_ Soil Horizons Reaction was determined in a soil-water paste (1:1 volume ratio) with a Beckmn glass electrode pH meter. Mechanical immes 93 _Sgil Horizons Mechanical analyses were made by the pipette method (Kilmer and Alexander, 19149) for the (50 micron particles and by dry sieving in the case of the sands. Determinations were made in duplicate. A 10.0 gram sample was placed in a tall 600 m1 beaker and the organic mtter in it digested with 30% H202 (technical grade) and heating. it the end of the digestion, while the suspensions were still hot, 1! H01 was added to the samples containing carbonates until further additions induced no reactions. The samples were then filtered through a 9 cm Buchner funnel with suction, using a hard (Whatman No. 50) filter paper. The residues in the funnels were washed several times to remove chloride and then transferred to shaker bottles. The suspensions in the shaker bottles were titrated with 0.1.1! NaOH to the phenolphthalein endpoint (pH 9) and then shaken for 21; hours in a reciprocating shaker. The dispersed samples were then washed through a BOO-mesh sieve. The sand collected in the sieve was oven-dried, fractionated by shaking in a nest of sieves on a mechanical shaker for 15 minutes, and then the resultant fractions weighed. The silt and clay passing through the BOO-mesh sieve into a sedimentation cylinder was diluted with distilled water up to a known volume and allowed to come to room temperature (constantly 20°C). The suspension *Dfleminations nude by Soil Testing laboratory, Soil Science Departmnt, Michigan State University. 118 was then transferred to a 2-liter beaker and while stirring, 25 ml aliquots were removed with a pipette, transferred to weighing bottles, oven-dried, and then weighed. The remaining suspension in the beaker was transferred back to the sedimentation cylinder which was then shaken end-cver-end (to disperse the sediments evenly throughout). The suspension was then allowed to stand for the lengths of time (calculated on the basis of Stoke's law) necessary for particles of a certain size and density to pass a given point in the suspension. At these calculated times, aliquots were taken with a pipette as before, oven-dried and then weighed. CHAPTER 10. RESULTS AND DISCUSSION Pollen m 9!. Three Lakes Egg The results of the pollen grain counts are shown in Table 6 and illustrated graphically in Figure 20. The data indicate that the beginning of pollen accumulation in Three lakes Bog was dining the Jack pine period suggested by Zumberge and Potzger (1956). If Breecher and Farrand's proposed chronology of lake events is correct, the bog could not have been in existence prior to about 10,000 years B.P., since the area was immdated by the Sub- Duluth deluge at approaintely that time. According to Fries (1962), this period was characterized by decreasing spruce and increasing pine (especially Jack pine) with pine dominance in existence by 9.150 2 130 years B.P. Evidence that pollen accumulation in Three lakes Bog began at the beginning of the pine period and not later is derived from the data of Iilson and Webster (19h2) which indicates that the initial doninnoe, the subsequent decline and resumption of dominance by Jack pine pollen occurred at the beginning of the pine period in north central Iiscomin. Fries' data indicate that Jack pine defiance lasteduntilabout 7300:qu B.P. innortheasternlflnnesota with other pines being dodmnt true that ti. until the late poet- glacial increase of spruce and fir occurred. Pollen from Three lakes Bog, howwver, indicates that Jack pine remined duinant until the late poet-glacial increase of spruce and fir occurred. This variance is probably related to the large area of sand soils in which Three Iahee Bog 'is located. Following the initial Jack and red pine dodname, white pines apparently began to succeed the other pines concurrent with an increase 119 FIG. 20 POLLEN DIAGRAM OF THREE LAKES BOG {0 'b' h I I I O. ¢)z~ 4 O O 00 «0‘9, 0 k .- 95‘") ~97 0 ~ W’II l'ce ' I 229 ”9 I I Is )3 '. o. e O *. C? u" - - I - I D * 5,? Q" w’ s 6’ — - — C _O _- - O — O 33Mbgj9 #30: 6" cf é‘ — — — — — -— JIM Ill 40’ k) .0 40 C) 20 I - -ee-k- - C - - e e I e eeII . O c '0‘. I I Il..lll . _ a II II I - I I I I - I I IceelIlIl I - ' I I I l IJ J' I II I e. - I I I I .l O O C) O C 0' 9 = e- 2: O 0 TREE pOLLEN 20 t) O O 20 40 60 80 20 TOTAL pERCENTAGE 0‘ 0 m.0 0 m.0 m.0 04 m.« 03 0.m 0 0 a...“ 5.3 «.3 0.2. 0m 0 m.0 0 0A 04 04 0.~ 0.0a m.» 04 .3 :4 0.2 0.mm mi. .2 0 0 0 m.0 m.0 m.0 0.... 0.0 0.“ m.0 0 04 men” 0.1.0 m.00 on 0 0 0 m4 0 0 04 0.2 m.m 04 0 0 0.3 «.mm m5. an 0 0 0 0 0.a 0 m.m m.~.n m.m m.0 0 a..." 0.3 WE 0.00 an 0 m0 0 m.0 m.~ m0 mA 0.3 0.0 m0 0 «A d? «.9 m.mm 0m 0 0 0 m2” 0...” 0 m.~ m.0~ 0.3” m.m m.m m.~ mfia 93 0.3 00 0 0 0 0 0.m 04 m0 m.0~ m.0.n 0.: 0.~ 0..” 0:: .18 0:3 mm m 0 m0 0 ms. 0.~ m0 0.m mean 0.0a 0.0 ~.~ 04 0.0 0:: 0.5 S 0 o 0 04 0.n 0 o.m. 0.5 m; m.m m.~ m.0 9n 0.0.." 0.5 m 0 0 m.0 0.0 m.0 04 m.m mam 0.m~ 0.m 50 Tu 0.~ «.3 m6” at m.0 0 m.0 man 0 m.0 0.m 0.0m men an 0.m m.~ 0 0.3 m.0~ m 0 0 0 m0 0 m.0 m.0 mAm 0.0a 1.0 0.0 04 min 0.3 0.~m N m.0 0 0 m...” 0 0.~ m.m 0.m~ 0.0." 02m 0 9m 0.m 0.0m m.~.m 0 n34 soothes bosons one: nooom a n8 nohm noose: a 095% eaten 3E 88 non .0300 than swarm 3a 3018 3.33 a 80.00 3.- MA .SE 8m mg as 2H mmgzmummm zaom an. .0 mafia 1213 0000000000 “} c: 0 00000000 0...” 0000 m.0 0000 0000 m.0 0.a m.0 m.0 m.0 m.0 o 0.~ m.m men 0.H 0.H m.0 m.~ m.H 0.H 0.H m.a moo m.a o..." 0 0.~ 0.4 0.0 0.H m.~ m.0 0.H 0.a m.a 0.“ m.~ m.n 0.0 0.n 0.m m.m 0.0a m.0 0.0 m.a m.: 0.0 m.m 0.m 0.m m.m 0.m 0.» m.a m.m m.aa m.0 m.0 m.H mvo m.0 m.a m.m 0.0 0.m m.n m.0 m.0 0...... m.0 m.~ m.0 m.0 00000 m.0 m.: 0...” 0.H 0.H n.a 0000000000 o.a h...” m.n «.4 «.0 m.m ~.H n.n .3 m.m 0.H n.0a 0.am «.mn a.m~ 0.H~ 0.0a a.m~ m.aa «.0 n.0w. >.ma menu H.o «.mn 0.4 «.mm n.a0 «.0n 0.ma H.0m 0.~0 m.m0 m.ma H.0m a.n0 n.ma «.0m 0.H0 0.:m 0.00 0.:0 n.aa m.mn m.mo m.sa mvam m.m0 0.00 0.Hm m.m0 m.m0 0.m0 m.ac m.0n mwao m.00 0.Ho n.00 0.0a aw." HNH 3H SH MHH 8H «OH 8: 00 as so «0 mm 122 in other genera such as m, Quercus and _Ulms;. Subsequent to a short period of white pine prevalence, however, the reverse of natural succes- sion seems to have occurred (i.e. white pins to red and Jack pine). This trend points to the possibility of a drying climate which would affect soil moisture condition relatively more on slightly developed sand soils than on medium—textured soils. Although this trend may have been brought about by regioml factors, it is likely significant that Green Bay was completely dry during lake Fayette and lake Chippewa tiles, thus the southerly winds characteristic of May through October at the northead of GreenBaywere likelyconsiderablywarneranddriertlnn previously or subsequently. This secondary dodmnce of Jack and red pine pollen persists fra the 110 to the hO-inch level where once again succession toward white pine is indicated. From the Ito-flinch level upward, natural succession under mister conditions my be indicated by the increase in hemlock, birch and beech pollen. The slightly later increase in spruce and fir begirmingatthe ZS-inchlevelistypicalofanusherofbogs in northern fisconsin, northern Minnesota and Isle Royals. This trend nrks the end of the 'hypsithernl' interval (Fries, 1962) which is cited at about 2000 years B.P. (Deevcy and Flint, 1957). mister conditions were likely brought about by the rise of water levels to the Nipissing level while subsequent cooler conditions with a con- current drop of lake levels could have been induced by an increased denimnce of dry polar air in winter. From the lO-inch level upwards, no olintic imlieatiom are at- tached to the fluctmtiou of percentages since lumberim activities my have influenced then. In comparing the pollen percentages at the 10-inch level with the pro-lumbering forest composition around the bog, 123 it Moons obvious that .1801: and/or red pine pollen overrepresents the local abundance of these species since about 17% of the pollen is Jack and/orredpinewithno evidence of Jackorredpine treeswithini-mile of the bog. Irite pine pollen percentages, however, do not seen to be much out of line with the relative abundance of that species in the local area. ml: and elm pollen percentages likewise do not represent local trees. Red nple trees in the vicinity of the bog are definitely underrepresented by the pollen, however. These relationships agree with the findings of Benninghoff (1960) and winch and Potsger (191:3). The presence of hickory and butternut or white walnut (M cinerea) pollen throughout most of the profile probably represents "long-range drift." The unusual occurrence (compared to northern Iisconein bogs) of both of these pollen types in the upper foot of the profile needs erplantion. The nearest known source area for hickory pollen today is60to70silessoutlwestchhree IakssBoginDichnsonCounty, Michigan (conversation with an unidentified forester in the kichipn Department of Conservation). The nearest known butternut trees are on StoningtonPenimula, sone25to30nilessouthof1'hree1akes Bog (persoml observation and correspondence with J. 0. Veatch). House plantings at closer locations is perhaps possible but no such obser- vations are onrecordas fares thewriterisaware norwere any observed by the writer. The fact tint Valders ice did not extend into the high country comment of Dickinson County (Hough, 1958 3 Thu-cites, 191:3) In! indirectly account for the occurreme of the call percentages of oak, ole, beamed, hickory and butternut pollen present at the lower levels of the bog profile. Curtis (1959) cites evidence indicating that 12h Valdsrs ice did not exert a refrigerating effect capable of completely elinimting oak and associated thernophilous species from the surrounding areas. 131 this event, post-glacial cliseral and successional ctnnges nest have been consistently nore advanced in the peri-Valders area of the Upper Peninsula thn in the Delta-Alger County area, not only because of the tine factor but because of the warmer, more continental growing season in the peri-Valders area of western Upper Michigan and adjacent Iisconsin. my and June tenperatm-es at Iron Mountain (Dickinson County) average 34; degrees higher than those of the Deltaqllger area, for instance. The interpretations presented above are subject to additioml in- accuracy resulting fron the possibility that peat accunulation rates and absolute pollen rain have varied greatly during the past. Certain pollen grains such as those of Popgus do not preserve well and thus the post-glacial ilportance of that germs is difficult to evaluate. Sons pollen profiles in Mimsota show high percentages of P9213 pollen within the spruce sons and the supraadjacent transitional sone (Wright, 1961;). making aspen, however, grows into drier clintes than the conifers (Spurr, 1961;) and thus post-glacial Ilinnesota conditions nay have been nore favorable for their relative abundance than post-glacial Upper Peninsula conditions. Correspondence of surface pollen percentages to sample plot forest cowposition (% of total basal area) also night not agree closely because of the possible lack of representativensss of the sampling point with respect to the total stand. ram A2212: The foliar analyses by plot and species are shown in Table 7. The 125 m2 ‘3 mm 3w .3” m. om." 2.0 0m.0 an. 3.0 $4 and .e 8.. 839m .8 4: ~30 0m m4.“ 0.» 4m 3m 0d0 3.0 mm". 2.0 3.0 8.8 .m 3.- 833 H0 .2 0.0 S 0.0m 3: 3A .30 3.0 2.0 min. 0.3 94 58m .4 is. 33 and «0 0m 0.: 40 mi: 0.0 2. a 3.0 34 d3. E..0 SA and! .0 ea.- 33 £5 «m a «.0 mun 0.5 0.2 0.: on 3.0 «.3 an. 00.0 mm.“ n23 .H 3.. 836m am man 4.... 3 0. mm 0.0 2. .RN 3.0 3.0 mam. 00.0 2.4 hood—om 8.. 838m .8 04 a... an .3: 4.0 n. S” 3.0 $0 «0“. 00.0 R4 .10 .m use. atoning 9 0m 04 on 0.3 «3 .3 «N 3.0 3.0 03. 0m.0 34 and .p on... p.833 «4 mm «.0 an mad 0.0 mu 3 3.0 F.210 mm". mm.0 «to «as .m use. stomach N4 00 0.0 .3 «.8 0A 3 nm 3.0 48.0 awn. 3.0 34 an. .m 3.. daemon .2 an IE IS IE an IE IS a u a u u .383 25. den .3 38 J a: nu m 8 on a. w. we a u a 995 do» «3.8 3a a fig .8 Ion—”Hugo go .5 "Ha 126 pioneer conifers (pines and balsam fir) have lower contents of N, K, Mg, P, and B tbn do the northern hardIoods, except that beech has relatively low I arxl lg contents. Sugar nple and beech have high contents of 1m colpared to all other species analysed. In the case of calcium, balsam fir foliage from a calcareom site Ins a content considerably greater the that of the pines on a calcareous site. Regardless of exchangeable soil calciun qmntities, the pines in this study contain lower amounts of foliar oalciun and nolybdenun than an of the other trees amlysed. Foliar conposition differences due to site are apparent in the case ofredandwhite pine. We andaluninun contents arenuch greater in the foliage from the Rubicon site (Cl) than from the Eastpcrt site (AZ); in addition, potassiun, iron, copper and boron contents are somewhat higher in 01 pins foliage the in A2 pine foliage. lack of response to site see. evident in the case of red pine foliar ealciun, phosphorus and ngnesiun. Foliage samples from both red pine sites (A2 and Cl) contain identical anounts despite wide differences inavailable Pand exchangeable Ca andlg inths soils. Definite differences between species are indicated where two or sorekinds oftreeswere saspledonthesansoiltype. Theredoak sasple contained decidedly higher concentrations of all the elenents except I, Zn and Al when compared with the pines. The white pine samples contained more N, Ca and Fe thn the red pine samples with the difference tending to be greater on the better developed Podsol. On the otherharnl, theredpinefoliage contaimddecidedlvmrennthn did the white pine foliage 3 the ratio between the two species was similar on both sites, averaging about 2.16/1. The yellow birch ealple contsdnd decidedly higher concentrations of Ca, Mg, Fe, Cu, 127 B, Zn, art! 110 than did the hemlock sample; mater amounts of N and lln in the yellow birch foliage are also indicated, but to a lesser degree. The hellock foliage, however, has concentrations of Al amounting to twice tint of the yellow birch foliage. The sugar maple foliage sample contains considerably lore N, P, Ca and In than does the beech sample, whereas the beech foliage apparently has higher concentrations of K, Fe, 011, Zn and Al. Similarities between site-crates also seen to be present. Foliar (x n11... of the two pines and red oak are quit. similar on Eastpor‘t sand. Thepines have similarvalues of KandPonEastportsarriand Rubicon sand. The yellow birch and hemlock samples contained similar concentrations of potassiun and phosphorus. Cornerning sugar nnpls and beech foliage, only llg seen to be similar, but both the sugar neple and the beech samples contained concentrations of Mn that were at least twice as high as those of the other couples. The foliar calcium values for pines, hsulock and beech in this stun are low but are within the ranges reported by others, Tables 1 and 2. The 3522 of percentages for the various species is essentially the sauce that of Bard for hardwoods and bollockandthe saneas that of Plice for fir and pines. Poliar potassium values for white pine and balsas fir foliages are very similar to those of Gerloff gt 33.. reported in Table 2. For hemlock and beech, potassiun values are sinilar to those of Bard listed in Table 2. For other northern hardwoods, potassiun values are higher than those ofCerloffgtg. butlowerthanthose ofBard. Balsanfirhadths lowest foliar K in this stew and in that of Chandler. Foliar phosphorus values for the conifers are siwilar to those found by Gerloff gt _a_l_... 128 Values for hardwoods are similar to or higher than those of Bard. Bard's chta indicate that, for most species, foliar P increases with depth to 11.. This trend is most obvious in the case of sugar maple but also seem to be quite consistent in red oak foliage. From this trend it night be inferred that more acid sites than those studied by Bard would give rise to still higher values of foliar P in the case of some species. Therefore the relatively high values of foliar P for sugar naple, beech and yellow birch reported here my be largely related to the soil acidity. Foliar N values in this study are sinilar to or somewhat lower than those of Gerloff _s_t_ _a_l_. but the values for hardwaods (except beech) are sinilar to those given by Bard for nature foliage. Values are within the ranges reported in individual studies ercept for the high value for henlockwhich ny be due to sampling inadequacy or to the nature of the site. Orders of percentages between species compare favorably with the data of Bard in the case of hardwoods (except for beech) and with Chandler in the case of pines and fir. Better correspondence might have been obtained between the data collected in this study and that collected in others had sites been similar, if there had been replications in this study and if the salpling procedures in all the studies involved had been standardized (m Ira-er and Koalowski, 1960). 'A' site conponents seen to be characterised by having relatively low concentrations of foliar K, P and Mn. Perhaps their status as pioneer trees on relatively unweathered calcareous sands is dependent on their low requirements for these three elements. For instance, white spruce, which is a frequent conponsnt on these types of sites, was found to have a low foliar deficiency level of potassium (Heiberg 129 and Ihite, 1951). The pioneer pines, however, undoubtedly cycle much less Ca than do fir and spruce on these calcareous sands. On the weakly and weakly to moderately developed Podzol sites with calcareous 'U' horizons (Sites El. and B2), hemlock is conspicuous with respect to its low foliar calcim content and its extremely high foliar aluwinun content. On Site Bl, hemlock and yellow birch are site-mates and have similar concentrations of foliar K and P. The similarities end there, however, with the consequence that these two species nest likely do not affect the upper soil horizons in a like manner. Specifically, yellow birch probably tends to raise the percent base saturation while hemlock tends to lower it. Supr mple and beech occur together on Site B2, and their foliage seem to be similar with respect to magnesium and high concentrations of wanganese found in then both. In this case, perhaps similar tolerances to high soil nnganese is important. Wilde (1958), for instance, nentiom that high contents of soluble unganese in Podzols seen to linit root penetration. If the high concentration of foliar P in the sum ample sample is representative, the presence of this species should have an appreciable effect on the chemical and biological properties of the upper soil horiaom. During the course of natural forest succession and soil development at these sites, certain trends in nineral cycling seen to be indicated. Foliage of pioneer balsam fir stands bring considerable quantities of calciun into the surface horizons of the soil. Should pines be the pioneer forests, however, much snller awounts of calciun are brought to the surface along with only small aaounts of the other ole-ants as well. The data point to the possibility that upon the continued ,‘ 130 dominance of the pines, strongly acid, weakly-developed Podzols eventually fern with the consequence that greatly increased anounts of Al and Mn are brought to the surface. it this stage white pine tends to bring considerably acre N, Ca, Mg, Fe, Cu and Mo into circulation than does red pine, while the latter species cycles much acre In than does white pine. is in the pioneer stage, the amounts of K and P cycled by each species are similar, with the concentrations of P being no greater than in the earlier stage of succession. If, however, succession to nixed stands of white pine, hemlock and yellow birch takes place prior to deep leaching of the carbomtes, weakly developed Podzols (such as that at Site Bl) my still fern, but the mineral cycling pattern will probably differ considerably from tint of the pine-dominated, weakly-developed Pcdsol sites (such as Site 01). Golpared to red pine foliage from the carbomte-containing sends the foliages of heelock and yellow birch contain concentrations ofN, Ga, FeandMothatareat leasttwiceas high, andPandB concentrations which are somewlnt higher. Cycling of aluminum by heslock is intense. Succession to sugar mple and beech involves the introduction of foliage with high concentrations of K (beech) or P (sugar nple) and In (both species), thus subsequent leaching lessee of these elements ny be thereby reduced. Although red oak is not necessarily a constituent of the sores leading up to the sugar maple-beech slim, this oak does occur through- out the Podsol Region of Michigan, and is mnerically dasimnt in some mtural stands and on new cut-over pine-oak sites. It is associated withredandwhite pins as wellas northerahardwoods. The red oak I. I. 131 foliage from Site 13 indicates that on very weakly developed Podzol sands, this species cycles concentrations of potassium similar to those of the pines. Phosphorus concentratiom are intermediate between those of the pines and those of the northern hardwoods, while nitrogen, calciua and ngnesiun concentrations are more similar to those of the northern hardwoods. Acid fall. leaching under pines should be quite severe since the forest floor is very strongly or extremely acid prior to September and the foliage indicates that subsequent increments of litter will not be rich in bases either. In northern hardwood stands, however, the flood of relatively basic leaves added to the forest floor in October should reduce the potentiality for subsequent acid leaching by forest floor constituents. Spring leaching, however, say be comiderable due to the increasing release of tydrogen ions (in the process of root respi- ration) and the replacenent of basic cations, some of which say not be tahenupbyplant roots andnythusbeleechedfroethe ecosystem in; m Fomtion Based on the theory of nor humus fornticn developed by Handley (1951:) and Davies 33 _._1_. (1960)* 1+. see-s likely that nitrogenom constituents in tln nesophyll tissues of dying pine, healcck, and beech foliage becoae stabilized by polyphenols. Since pclyeerisaticn of polyphenols is favored by base-rich condition, and polymerisation beyond a certain aelecular sise precludes aw tanning action by “The theory proposes that pclyphenols in dying leaves stabilise the proteim in the eesoptyll tissues. These stabilised proteins protect the sesopm‘ll cell walls from deco-position and together theyforaanalorpheus residue lying on the surface of the mineral soil. 132 polyphenols (Coulson 31. 21., 1960), it is probable that the low calcium and ngnesinn content and/or the low antacid buffering capacity of the foliage of the above species is conducive to the tanning process. This wpothesis seems logical since the chemical characteristic that most clearly separates the null horizom from the nor horizons in this study is the exchangeable c. (flue): organic utter (%) ratio which is above 200 in all the Vh horizons and below 100 in all the Oh horizons. S32 Microflora Studies Direct microsc0pic counts of nicrcflora in the Oh horizon of the Site Bl soil. and in the Vhl horizon of the Site Dl soil are tabulated in hbles 8 ani 9. These horizons represent the extremes of nor and null bums types found in this study. Counts of bacterial cells didnot reveals mimggaggfsoil in the null husms horizon but there were twice as new cells p_e_r_ m 95 95.89.13. git}; in the null. horizon as in the nor horizon. The null horizon contained vastly greater amounts of actinomcete filaments; and per gran of organic matter, this horizon contained 25 times as much as the nor (h horizon. Amounts of unstained fungal mphae were greater in the nor 0h horizon than in the null Vhl horizon. Again, however, amounts of stained and unstained twphae per gran of organic utter were greater in the null m. The presence of greater amounts of actinoswcete filaments in the Vhl horizon is in accordance with previous studies which indicate that environments having a pH greater than about 5.0 are more favorable than those having a lower value (Alexander, 1961). High numbers of bacterial cells/g of soil in the nor Oh horizon are likely the result of the bid: 83.8 n m 535 :3 n ma 3.3335 n no 358833 .85 .. 55 5 z a £85m u 8.88 55 «no.8 u £38. .983... be»! chaise u. mmoa 8o. «.4: «a. an“ 5.2. 58. 25 «.8 35 u .. {8.2 .Smsmw an... 55 2m 8. 3: an 5.3 18 85. we. 18 an.“ n .. 78:5 .22 33.3 “do. e935 Aw\.ac .oev Ba 80. 2.5 8H 55m m5." 08. 5mm. m.m.n 3.." .. 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N...— mum .. ... a .. .. .. .. u .. .. .. a .... 3 .898 c.9835 3.335 Edam salsa .38 .353 BE 3.33 $385 aHm Em. «noun—”8m no man. 2H ”Egg .3 Eco amooag Han on ands mh<\§o.n u m ode-Pd «an n ma .m5\hm4n u 8 35882.5 «8.4 u 55 5 a a $3.9 . 83:8 use... nae»... £35m .- 8.8. .fiefifl. 7.3.23 8mm 85 33 ms 45m m.4m 85 4.3 a5m 4.4« u .. .35 u .92 8.er Ads. 3 .32. .n.o SH 4.«« fig $3 at: n5." 85 «m.« «m5 55 u .. {.84 8.25 78.65 duos pedal m5m n5H R5 m.4« 44.m $5 85 3..." 85 04.." .. .. {.23 8.5m 2.4. m8. H8. 48. 58. 48. 85 48. 58. 58. .. u 3.853 55 3.5.3» efloofic «8. do. 48. a8. o8. So. 85 a8. 58. 58. .. .. HS. 55 mung?» 0.50 58. . m8. «8. m8. 8. .8. 85 m8. 8o. 8. .. .. do. .3... {Indus 304 m3 5% 4mm... «on mm» 85 mes an HR am .. 55 3:3... So fimn two m4m an" 4.8 85 55 o.«4 454 5.8 .. d8 55 £33.: 2.« 4.3 55m ems nae 954 85 a5 «.2 05a QR .. 3.. p.8- £28.: I .. .. u .. .. .. u u .. .. «S a «A .25 noon .. .. .. .. .. .. .. .. .. u .. 53 a a..." a... 55 u .8 u .. .. u .. ... .. .. .. a .. sea a on. do. 3... «an E 3i. W4 E E QTEIIE uuuuuuuuuuuuu 3 88c noes—:5 338s e33... 3 .58 .55 Hanna 38353.4 3335 *HQ En «Ramon a nun. 2H 9380s .8 menace H.338”! an em and“... 1.35 percentage of organic matter in that horizon; on the other hand, the relatively low numbers per gram of organic matter is possibly related to the greater decomposition resistance of nor organic matter as com- pared to that of null. Although anti-bacterial substances may be partly responsible for this low density, the lower N content of the organic utter in the Oh horizon could also be of significance. The relatively high content of unstained fragments of fungal nyphae (representing dead fungi according to Jones and Molliaon) in the nor a: horizon agrees with the findings of Ronell (1935) and statements nds by Wilde (1960). The fact that the volumes of these fragments comprise as much of the weight of the organic mtter in the null. m as in the nor a: is due to larger quantities of mphae of large diameter in the Vhl horizon. This hyphal diameter difference points to a difference in specific composition of fungal populations in the tie horizons. Comparisons of hyphae volumes per unit volume of organic ntter, however, cannot be made from the data available. Despite the possibility that the results of this study may have only limited applicability, it seems worthy to mention the possible significance of relatively high snounts of actinomete filaments. Alexander (1961) states that mess organisms are relatively scarce during the initial stages of plant residue decomposition but becme more prominent later when nutrient levels are lower and competition from other organism is less. Their competitive advantage in the latter stages of decomposition probably stems fro. the fact that they can utilize such carbon sources as cellulose and chitin. Since chitin is a cell wall constituent of may fungi, the larger surface area of unstained typhae in the Vhl horizon might be a factor in promoting 136 the larger number of actinonycetes in that horizon. Actinonycetes have also been found to have the ability to oxidize polyphenols to quinones through their production of phenoloxidases (Kononova, 1960). Kononova further states that a number of studies have shown that the condensation of the quinones thus formed produces complex, dark-colored humic substances. harness-s wwwgmuu en Total carbon concentrations were invariably highest in the surface horizons. When converted to percent organic matter, these surface horizon values range fron 14.87% in the V111 horizon of Profile A1 to 51.0h% in the Oh horizon of Profile C2 (see Appendix). Pr0file distributions of percent organic matter and total nitrogen are shown graphically in Figure 21. These graphs show that a secondary peak of organic natter occurs in the subsoil of all profiles with an E horizon. This peak is most pronounced in the lhib horizons of Profiles 02 and D1 (the only nderately developed Podzols), a result expected since the dark colors of these horizons indicate a relatively high organic matter content. However, the organic matter percentage is apparently not the only factor contributing to dark illuvial horizons. For imtance, the lhib horizons of Profiles B1 and B2 contain 0.78% and 0.79% organic nutter, re- spectively. The Munsell color notation of Bl-Ihib, however, is 51R MB while that of B2-lhib is SIR 3/6. The upper illuvial horizons of Profiles Cl and D1 contain 1.13% and 1.16% organic matter, respectively, while their colors are moderate brown (7.5m 14/14) and dark grayish brown (513 2/2), respectively. Conversion of 0.11. percentages to the estimated number of pounds per acre results in the following values for the illuvial zones: (INCHES) DEPTHS and HORIZONS 137 FIG. 2I PROFILE DISTRIBUTIONS OF ORGANIC MATTER AND NITROGEN O 8 Percent Organic Matter 0 = Percent Total Nitrogen %0M.0l2-345678 l00l23456789l0 0 ——e ’20 4O . Site '8 2 60 Site Al Weakly-Mod. Developed Regosol Podzol , lAleJlLllllJLlAlAl legjlllzlJLnlkl-zls ‘70 N o 5 LO 0 5 LO Q“ t v v v v r ' r ' ' t w I I I V O - a 20 40 " Site AZ . 30 Very Weakly Developed Site Cl Podzol Weakly Developed Podzol 40 P Site A3 60 Very Weakly Developed site 02 P°d‘°' Mod. Developed Podzol Qh "IN 20 40 3° 3m Bl sm on .Weakly Developed Podzol Mod. Developed Podzol 138 Profile PoundsLilluvial zone /acre n .- 12 55% A3 6361: B1 h2,120 B2 19,628 01 141, 762 02 5h,356 D1 57, 753 These values indicate that illnviation of organic utter has been taking place at a faster rate and/or for a longer period of the in Profile Bl tlnn in Profile 32 although the soils are chronologically ofthesaneage. ThesauthingistrueofProfileCZasconpared Iith Profile Cl. This difference in degree of development may be related to the length of tine chring which the sites have been characterized by nor humus layers and hemlock-hardened forests. §_o_i_l_ 9353313 m Fractiomtion The percent humic acid of the illuvial horizons from Profiles B2 and D1 (hardwood null hunus sites) exceeded the percent organic matter, thus pointing to the possibility that clay as also present in the hustle acid fraction obtained. Fulvic acid was present in all horizons con- taining organic utter, the brainsh orange or brownish yellow colors resesblim very closely the color of concentrated crenic acid described later. The results of several studies indicate that hardwm lignim contain spiral groups whereas conifer lignine do not (see Kononova, 1961). R. I. Morrison (1958) used this distinction to identify the source of organic utter in sons Scottish soils and posts. The thereo- decouposition technique used in the present study suggested that urinal groups were absent even in 100% hardwood soils. Guaiacol 139 and phenols were present in the humic acid fraction from both more and nulls (Bl-0h, Bl-Vh, B2-Vh2, D1-Vh2) and in the Ihib of Profile B2 (in this soil, the V112 grades directly into the Ihib). In the D1 soil, no similar components could be found in the Ihib2 (distinct E horizon separates V'h from I horizons), suggesting tint illuvial humus has a different composition than Vh bums. llthough the origin of the illuvial organic matter could not be determined as anticipated, it is clear that the organic Hatter of the Oh and Vh horizons studied contains mterials derived from lignin poly-ere; thus the nan “ligno—mcelial nor” (Wilde, 1958) seem quite appropriate for the minus layer of Profile B1. The presence of illuvial horizon tunic acid does not necessarily imply movement of this fraction since hmnic acid can be formed from fulvic acid in the presence of iron iom in an acid mediun (Sheffer and Ulrich, 1960). Martin (1960) states that the relative anounts of these two organic matter fractions seens to be controlled by the amount of ionic L1 present; his studies indicate that ionic Al is capable of flocculating Podzol humus at pH values comonly found in Podzol soils. m 529 Production _o_f_ §2L1 Horizons At the onset of the incubation and leaching stuw, obvious anounts of acidic, yellow organic utter were noted in some leactntes while others contained none. Berzelius applied the torn 'crenic acid“ to such acidic, water-extractable organic nterials. Crenic acid, or components of this fraction, hve frequently been referred to as important metal- couplezing agents in the fomtion of Podzols. The pattern of crenic acid production for each sample is shown graphically in Figure 22. 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Calculations of crop nutrients per acre was made on the basis of the percent composition per species with interpolations and extrapolations being made for species for which no total. tree percent canposition data was available. The interpolations and extrapolations were made by com- paring published data on foliar composition of the species involved with those for which total tree composition data were available. *An average DBH of 13.5 inches was used since the basal area of that diameter is approximately 1 square foot. Thus, if the basal area per acre or I species 1:: 80 sqmre feet the total weight of a 13.5 inch DBH tree can be nultiplied by 80 to arrive at the stand weight per acre of that species. 235 APPENDIX V. 5011 SERIES DESCRIPTIOI‘B 236 237 1 . :k caries consists of well droixci workiy 0000*0301 yo no; :02...s l"uf“fi r5 '; ,;:olic soils which developed in acid fine s ads and ssnls on dunes near t a L 11. They have faint A2 horizons and uszzally have thin Bzir horizo.1s of sli~htly or chroma than either the A2 or the underlying C horizon. They have less dist1nct .cizonation and lighter colored BZir horizons than the Rubicon and Vilss soils, and are formed in finer sands. The Deer Park soils differ from the Grayling and Omega smile I having more distinct A2 horizons. Deer Park soils are devel pad in sands in the 1‘1: 1d of t'.e mec'icm sand size and in fine sands; while Grcyling and Omega“ are sloped in sauce in the coarse and of the medium sand size a11d in coarse so ds Der: -;; soils include profiles with 221: hor's as comparable to those of G1 ayling an d “a at the most '”'ongly developed and of their range. Deer Park soils also a)»- :‘1Lc soils without identifisole E21: horizons, which the 6:9 cgling and Omega soils #2:. The? hive loss df.stinct horiso:.ation than the Eastport soils and are medium at;‘ 72:23.0 of so moi feet, while Eastport coils are sligltly acid tozildly elic.ire r“: so? a sni overlie calcareous or no: crately allcsline sands at depths of 25 :3 or 1 eco. Deer rsLR soils have much more weakly developed Podzol horizons ctr: 'géu " 5‘3 and are coarser in to"turo. roar Park soils are more sold; h.cv. lose I f : lax: the strctiiiznt. .cn of t1 3 Lhccota and retest: 13113. Ice: :r‘“ 'v cgcc'si a is the dune use“s alo~3 €12 61cc.t Lalca 31c c e 'gcd 1:14: y 1 0: cl purposes. Cflffisijlci. Deer Park sand. C~1"Vcry dark brown (lOYRZ/Z) to very dark gray (lOYRB/l); sand; single 5 structure; loose; contains considerable organic matter; 1/40100.h can of forest litter on surface; medium to strongly acid; clear smooth boundary. 1 to 3 inches thick. 104” Light bro"nish gray (171'R6/2); sand; single grain structure; los:c, mszy fin1c roots; medium to strongly acid; clear wavy boundc1y. 2 Lo 6 inches thick. 7 4230” Yellowish brown (lOYRS/é) to light yellouis h brOJn (2U 0'36l) sand; single grain structure; loose; common fine roots; medium to strongly acid; gradual wavy boundary. 12 to 25 inches thick. 20060"+ Pale bro"n (10"R/ 3); s and; s1’ngle gr sin 3 ructure; loose; few fire roots extend to 63 inches; mc'ium acid; no y feet thick. 5-".§5::1::1f‘°§111i Sand and fine sand types are recognized. The A2 horitt 0.1 es is as b: igi1t cs roan (lOYRS/3). Where the A2 horizons are thic303t, Lie son is on1"s lightly lower in c::1ro:n than t 1c 1311' 110112011 he? 01: it, (10-2.: only ‘*r. The BZir horizon is usually present, and may range up to 25 121cc " LZir horizon is never well expressed, and usually has values of 5 or g. cm usually ranges in thicl.css from 18 to 36 inches. Fine sand P'“Lro often ‘2- :’.n the unyer 3011101616 with the con’nnt of mod ium sands smuetincs increasing 1.. 013 1c. er to moist conch tions. l' . we ‘3 u ‘0 1. 1 .'_. , ‘ ;'-,9“': Ccic-c to seaop sand dunes. ..--..J.“..D slow to urdfun: '0 m D ’- '1 rs He L) :1"1_cjd Forrothilitv: Well drained, Sur: c runs” .. ... ' ~ - y “ .101111y is r0p1c. . I ~ . . "~. ... 1 V ,. 4 v . f. O _1‘ 1‘f\".". ... 6 4—‘ a...‘ 1‘r~.-‘r1’r.. 0" , a .1. \. s ’.. v - k.'. l . a. -- o..\.- \-o- ..nq . V. V o -~v\fi " l 'r p 7.-.‘ "— ’9‘, as,- -L- A - - — J ...- ;,,J-...La ‘ '~ ‘ . ; - I ..‘ ' --..-.‘ .. . . .., r‘. (1_ R, ‘ .‘ « 7-- h p . .. _ -.. < J 0 ~.' I d _ .-.. 0 fi' ' . . ‘ Q 3‘" '\ .t\.-, 1 , ‘ , o - — .} ‘.‘ 4— ‘J N". a“ " - .... . . ¢ov a .. ..x’. . v . - '. - ‘ _ ‘ : l 1 (~- ‘ VI. . .~ 0/ “ 1 1 . m" ‘2‘-" -.L - ~. ..a. ,, 4 \. .\. . I Q r. «- u; 1.. A 0 V f‘~ x..- » C’ ;J . . 1’ 0‘ .- x. f 1.... ...,9; . ...-g... tied 5- -.. '3‘! .. .‘1If u . . 11".”.‘Cy '2' L 0‘ , ’ 4. r. I ‘5“ G g up‘ A , vv‘ \ -\ 238 fl. VI. .. ~ . a ., . 4. a I :l.. ., ¢ . ....— _ . . v 5.. . . onlv . I ’ I ' Ia i. J.,). 1‘. \a _ “n. (A. u |. Of, ... 3 I, J .. .. v .. v .. , . . v .z‘ e . w ‘ . T ..Q .. «I; my . u f... s 1.- .. .‘ .xf _ t.) O n . . . ul- I“ :1; . , . L3.“ . ‘.'l¢‘-l‘.4:§-! ./ I I \ x r. n. - u. .A . fl‘ . ‘1 «ii:— \ . . A lo .v.. . “ r .. .I .. Q o u .. . o. . .2? n ‘ ‘,“ Ark ...! fl u. ‘ a v . v I s 7.54 a. ‘ xi H.» k .L '1 ... .- v. .‘o I. o “a... .. .1 D O, . . ‘ «v .1. wry: 0 \ 4, .. .. I 5 4 \ fl“ ‘ . u . « a ‘ . . ’1‘ . ., , \. . V11 ..T x .\ I LI, . 4 ..v s 1 D’nl‘ . 1.. v. « J. . .I a - ~75, 239 . 1 ; . K A ,' I ‘ . , \ A ‘ , . f r n, _ ' ‘p ' ‘ I . f I' s ; ' v ‘ / ‘ ‘ ‘ , ) ¢ . i ‘ 7‘ I .‘ . ‘ 1._ . VI ‘ ' ' ' \ " u a v " ‘I . '4 ‘ I x I ’ ‘ . . r ‘ ‘ r ' ,t‘ “ .1 ’N 5_ , , ‘ I A - , '. I u I '4 '1 . ., U AI'k "V ‘ m vr...,‘v.-, Awé v; .4 ‘ . ‘ 4 I ‘ r'l- ,r 'l _. ‘ . / ' C . ' . . V! ‘ r‘ . ‘ h x; 3: , ‘ 'L‘ ‘ .-‘ . ' at} g’ n 5 . L. I ‘ .11: I ' . . _ ~~ ~. ' . ' , 1 u - ' - ' "' ‘ ‘ I A J ‘ . \ ': «I ‘1’ \‘ ‘1', I j. ) ‘ ‘ 1" :- V l 2&0 5.} lflTA E ERIES The Sparta series comprises somewhat excessively drained.Regosols intergrading to the Urunizem group. They have developed in acid sandy parent material on level to gently sloping outwash plains and stream terraces under a grass cover. The parent material consists almost entirely of quartz sand and contains very few otheT minerals. Sparta soils occur in close association with the Plainfield and.Gotham series. They differ from Plainfisld soils in having a thicker and darker colored A horizon. Gotham soils have a slightly lighter colored A horizon. a thicker solum, and a B horizon with slight clay'accumnlationo lacking in the Sparta serissc Sparta soils have developed in 0081 'C so textured parent material, lack the B horizon nri.ha.ve a thinner solum than the Dakota soilsr. Hubbard soils have a B horizon; t12cter s l , and have dcvelopedf Tom mixed 1i “hologic material. Although Sparta soils are r tEer widely distribxtod their total acreage is relatively small and they make 1 ttle contribution to agricultureo 1: Sparta loaay fine sand - meadow 0 Q3 2 <2 )"3 r.» ;2y'dark gray (10?? 3/l) to very dark brown (101R 2/2) cany fine .. . - 1 I)" -, ._ 1:1“ (...) ..L f.. K -‘-1 f." . . a, PM .P n ,9 o "iv-'72 5 0'. 2:12:30 3‘ '! -;r :‘20 - _ .- {07- 1.2‘ C -. T-‘JxI: ::t;3~‘k J— 1-;2KJ ‘x :33," L’L‘ JV 2" 5. C, ’ ‘reo'. y .. n‘b‘AJ.) .r ’ .3:ch ‘21:. $1. a\. v) ‘ «i- a A1 .2, «:0 ... 3‘ j Q, \ a“ — .; .- u'Q—v A w~, . IU-‘v-p n '0 1.!) -_ u \ 44‘TQ‘90? ,g 1“ \. \I‘ Vat; J.L'3.; '44.; 1:»; o l..:21..4 U- L14. s CT #381 WC.‘.' ‘ bulludui o 120 ‘ 44.1.4 1;;3) . H .11... ? L2.» “Gr. 0 r ~ ‘m a» " I \ ,3 9~18“ Lcrr brown (10m 3;,) to vo ”y'd T1{ grayish bro":i (lOIR 3/2) lo Ty " fine sand.; vs ;-'y fiiie: weak g:anu;ar structure; vs TV fii3bls; Toots p1entiful;s+"cngly to medium acid; clear waxy boundary. 8 to 12 isohes thick. Cl 18n26” Yel].owish brown (1013 4/4) fine sand; single grain; loose; for plant Toots; very siTongly to s ongly acid; indISLinct boundazjo 8t to 12 inches thicE ft - r u o o q C? 23”? lellcwash.brown (1m MS/ ) to brownish yellow (lOYR 6/6) fine sane; single gain; loo so; we Mil stratified; strongly to medium acid beco2 12g more neatly nou trs .1 with depth. Se oral lest thick° 12:22: i: 9p;rffj:rws-20s° Color of Al hcrLzon ranges from black (lClR Z/l) to ‘Cif oarhgr';ir 2h 01 OJJ (lOfR 3/2)01ne 91 ran: as in thicE:ness from 8 to 2 incxes. Thin noncontanuous lenses or clayey