MIXED CONIFER and RED FIR FOREST STRUCTURE and USES in 1899 from the CENTRAL and NORTHERN SIERRA NEVADA, CALIFORNIA Author(S): Scott L

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MIXED CONIFER AND RED FIR FOREST STRUCTURE AND USES IN 1899 FROM THE CENTRAL AND NORTHERN SIERRA NEVADA, CALIFORNIA Author(s): Scott L. Stephens Source: Madroño, Vol. 47, No. 1 (JANUARY-MARCH 2000), pp. 43-52 Published by: California Botanical Society Stable URL: http://www.jstor.org/stable/41425343 Accessed: 24-06-2015 21:50 UTC

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This content downloaded from 169.229.32.36 on Wed, 24 Jun 2015 21:50:04 UTC All use subject to JSTOR Terms and Conditions Madroño, Vol. 47, No. 1, pp. 43-52, 2000

MIXED CONIFER AND RED FIR FOREST STRUCTURE AND USES IN 1899 FROM THE CENTRAL AND NORTHERN SIERRA NEVADA, CALIFORNIA

Scott L. Stephens1-2 Watural Resources Management Department, California PolytechnicState University,San Luis Obispo, CA 93407 2CurrentAddress: Division of Forest Sciences, Departmentof EnvironmentalScience, Policy, and Management; Universityof California,Berkeley, CA 94720-3114 stephens@ nature.berkeley.edu

Abstract

Historicaldata collected from five "average" mixed conifer stands, four large mixed conifer stands, andfour red fir stands from the central and northern Sierra Nevada by George Sudworth in 1899were analyzedto determine historic forest structure including diameter distributions, basal areas, and snag and livetree densities. The effects of early logging operations on standcomposition and structure is quantified by comparingcharacteristics of the trees that were harvested versus those unharvested in four mixed coniferstands. Average diameter atbreast height (DBH) was86 cm(34 inches)in the "average" mixed coniferstands, 110 cm (43 inches)in the large mixed conifer stands (this was equal to the average DBH of8 mixedconifer stands sampled by Sudworth in thesouthern Sierra Nevada), and 77 cm(30 inches) in thered fir stands for trees greater than 30.5 cm DBH. Shadeintolerant tree species dominated the "average"mixed conifer stands, shade intolerant, intermediate, andshade tolerant species were abundant inthe large mixed conifer stands, and Abies magnifica Andr. Murray dominated the red fir stands. Mean treedensity for the "average" mixed conifer, large mixed conifer, and red fir stands was 229 trees/ha, 235trees/ha, and 433 trees/ha, respectively. Average tree density was higher in Sudworths southern Sierra Nevadamixed conifer stands when compared to thecentral and northern Sierra Nevada. Snag density averaged5/ha in the large mixed conifer stands and 17.5/ha in the red fir stands. Early logging operations removedthe majority ofthe Pinus spp. and Pseudotsuga menziesii (Mirbel) France leaving large amounts of Calocedrusdecurrens (Torrey) Florin and Abies concolor (Gordon & Glend.)Lindley. Information fromthis study can assist in the characterization ofhistoric stand structure inthese forest types.

The absenceof firein the20th century and past Analysisof historicaldata have been done for harvestingoperations have modifiedthe structure the Stanislausand Lake Tahoe Forest Reserves and ecosystemprocesses in the coniferousforests (Sudworth1900), portionsof the northernSierra of theSierra Nevada. An increasein thedensity of Nevadaand the Transverse Ranges of southernCal- small shade toleranttrees has been producedin ifornia(McKelvey and Johnston1992), and por- manyforest types (Leopold et al. 1963; Hartesveldt tionsof the southernSierra Nevada (Stephensand and Harvey1967; Vankatand Major 1978; Parsons Elliott-Fisk1998). All ofthese studies discuss early and DeBendeetti 1979; Bonnicksenand Stone loggingoperations but no workhas been donethat 1982) and thisincrease has resultedin a decrease quantifiesthe effectsof earlylogging at the stand in forestsustainability (Weatherspoon and Skinner level, the amountof hardwoods van quantifies present 1996; Wagtendonk1996; Stephens 1998). historicallyin mixed coniferforests, determines Changesin climateover the last centaury may have historic densitiesand or differentiates also contributedto the in foreststructure snag sizes, changes betweenaverage and maturemixed conifer stands. (Millarand Woolfenden 1999). Early loggingoperations affected the composi- Historicaland prehistoricinformation on the structure size and tion and structureof Sierra Nevada forests,es- (density, distribution, species between1860 and 1950 and composition)of mixedconifer forests are relatively pecially (Laudenslayer rareand have been reviewedelsewhere Darr 1990). In 1899, approximately45 percentof they (Ste- thetrees harvested in Californiawere either Pinus phensand Elliott-Fisk1998). One of the methods thatcan be used to determineprehistoric forest ponderosaLaws (ponderosapine) or Pinus lam- structureis the analysisof data fromearly forest bertianaDouglas (sugarpine). Most earlylogging inventories.These data providequantitative infor- operationsin theSierra Nevada harvestedall trees mationon historicforest structure, however, the re- thatwere considered to be merchantableat thetime sultsfrom the analysescan be biased because the of theharvest (Laudenslayer and Darr 1990). methodsused to selectthe standswere frequently The viabilityof theCalifornia spotted owl ( Strix notrecorded (Stephens and Elliott-Fisk1998). occidentalisoccidentali s) is receivingmajor atten-

This content downloaded from 169.229.32.36 on Wed, 24 Jun 2015 21:50:04 UTC All use subject to JSTOR Terms and Conditions 44 MADROÑO [Vol.47 tionin California.The owl prefersto nestin mixed analyzedin thispaper withthe exceptionof one coniferforests with 80 percentof thenesting sites standlocated in a purePinus jeffreyi forest because occurringin thisforest type followed by 10 percent of no replicationin this foresttype. Exact stand in Abies magnificaAndr. Murray (red fir)and 7 locationsare not givenin the fieldnotebooks but percentin Pinusponderosa hardwood forests (Ver- referencesto rivers,mountains, and landmarksare neret al. 1992). The remaining3 percentof nests included(Sudworth 1899). occurin eastsidepine forestsand foothillriparian- Five "average" mixedconifer stands, four large hardwoodhabitats in the westernSierra Nevada mixedconifer stands, and fourred firstands were foothills(Verner et al. 1992). recordedin the 1899 fieldnotebooks (Sudworth The habitatrequirements of theCalifornia spot- 1899). Mixedconifer stand data were stratified into ted owl have been investigatedand it nestsin old- two classes (averageand large) whereasthis was growthforests with high canopy cover (Gutierrez not done in the southernSierra Nevada analysis et al. 1992). A relativelyhigh number of snagsand (Stephensand Elliott-Fisk1998) becauseno stands downlogs are also correlatedto thecurrent nesting were identifiedby Sudworthin his notebooksas sitesof theCalifornia spotted owl (Gutierrezet al. having"average" characteristics. 1992) butno prehistoricaldata existon the abun- Sudworthrecorded the species, diameter at breast danceof snagsor fuel loads in thisforest type mak- height(DBH), andnumber of 4.9 m (16 ft)logs for ing it difficultto describethe compositionof the each treegreater than 30.5 cm (12 inches)DBH prehistoricalhabitat. (one 28 cm DBH treewas recordedin a red fir The objectiveof thispaper is to analyzemixed stand).Each standwas sampledwith one 0.1 ha coniferand red firforest inventory data acquired (0.25 acres)plot. He recordednotes on regeneration by GeorgeSudworth in 1899 fromthe central and (estimateof densityby species, not a complete northernSierra Nevada to furtherour understanding seedling inventory), forest floor depth, and other of forestconditions and theirmanagement in the informationsuch as therevenue generated by early late 19thcentury. Analysis includes snag and live loggingoperations. He also frequentlycommented tree densities,basal areas, diameterdistributions, on theeffects of earlygrazing and burningon the and quantificationof the effectsof earlylogging SierraNevada and his commentsare summarized operationson standcomposition and structure. below. The followingvalues were calculatedby aver- Study Site and Methods agingall standdata foreach of the 3 foresttypes ("average" mixedconifer, large mixed conifer, red The historicdata analyzed in thispaper were ob- fir):number of snagsper hectare,snag basal area, tainedfrom the area of the centraland northern diameterand speciesof trees removed by earlylog- SierraNevada thatnow includesthe southern por- gingoperations, basal area per hectareby species, tionof the Tahoe NationalForest, the El Dorado numberof treesper hectareby species (density), NationalForest, and northernportion of the Stan- quadraticmean diameterby species,percent total islausNational Forest. basal areaby species,and percenttotal stocking by Mixed coniferand red firforests were surveyed species. in 1899by George B. Sudworthwhile employed by Standdata are summarizedand discussed,but a theUnited States Geological Survey. The purpose statisticalanalysis was notperformed. Selection of of thissurvey was to inventorythe forest reserves an appropriateanalysis method requires informa- ofthe Sierra Nevada. The originalunpublished field tion on samplingprocedures which are unknown notebooks(Sudworth 1899) werethe source of the forthis early forest inventory (Stephens and Elliott- inventorydata analyzedin thispaper. Fisk 1998). SierraNevada mixedconifer forests sampled by Sudworthwere composedof whitefir Abies con- Results color(Gordon & Glend.)Lindley (white fir), Abies magnifica,Pinus ponderosa, Pinis lambertiana, "Average"mixed conifer stands. The fivemixed Pinisjeffreyi Grev. and Balf (Jeffreypine), Calo- coniferstands denoted as "average" by George cedrusdecurrens (Torrey) Florin (incense cedar), Sudworthwere dominated by moderatesized trees Pseudotsugamenziesii (Mirbel) Franco (Douglas- of severalspecies. The averagequadratic mean di- fir),and Quercuskelloggii Newb. (California black ameterfor all treesover 30.5 cm DBH was 86 cm oak). The red firforests were composed of Abies (34 inches).Average tree density was 229 trees/ha magnifica, Pinus jeffreyi , Pinus montícola Douglas (92 trees/acre)(range 150-300 trees/ha).Average (westernwhite pine), Pinus contortaspp. murray- basal area was 130 m2/ha(558 ft2/acre)(range 94- ana (Grev.& Balf.) Critchf.(lodgepole pine), and 186 m2/ha).Table 1 summarizesall standcalcula- Tsuga mertensiana(Bong.) Carrière(mountain tionsfor the "average" mixedconifer stands. hemlock).Red firforests are widelydistributed and The largesttrees in the"average" mixed conifer theycan be foundon boththe west and east sides standswere Pinus lambertianawith an average of theSierra Nevada (Rundelet al. 1977). DBH of 108 cm (42 inches).The largestPinus lam- All standdata recorded by Sudworthin 1899 are bertianarecorded in the inventoryhad a DBH of

This content downloaded from 169.229.32.36 on Wed, 24 Jun 2015 21:50:04 UTC All use subject to JSTOR Terms and Conditions 2000] STEPHENS:MIXED CONIFER AND RED FIR FORESTSTRUCTURE AND USES IN 1899 45 Table 1. AverageCalculations of GeorgeSudworth's 5 "average"Mixed Conifer Stands in the Central and NorthernSierra Nevada in 1899(Standard Error). * Averagevalue for all stands. Basalarea DBH Percent (m2/ha) Trees/ha (cm) oftotal Percentof Tree [130]* [229]* [86]* basalarea trees/ha Abiesconcolor 5.3 6.0 105.7 4 3 (5.2) (6.0) (0) Calocedrusdecurrens 26.0 54.0 80.6 20 24 (6.6) (11.2) (10.5) Pinuslambertiana 8.8 12 107.8 7 5 (4.2) (7.4) (22.8) Pinusponderosa 56.6 106.0 83.9 43 46 (23.1) (37.0) (12.1) Pseudotsugamenziesii 30.9 38.0 101.2 24 16 (20.2) (24.6) (1.8) Quercuskelloggii 2.4 13.3 58.0 2 6 (L4) (3/7) (7/2)

152 cm (60 inches).Pinus ponderosa was themost fourstands were harvestedalong with88 percent commonspecies comprising46 percentof total of thePinus lambertianatrees (Table 3). The ma- stockingand 44 percentof totalbasal area (Table jorityof thewood harvestedfrom these stands was 1). fromPinus ponderosa trees with an averageof 64 Abiesconcolor was rarein thestands accounting m2/ha(275 ft2/acre)removed and this was 2.4 times foronly 3 percentof totalstocking and 4 percent greaterthan Pseudotsuga menziesii which was the oftotal basal area.Calocedrus decurrens and Pseu- nextmost common species harvested. The amount dotsugamenziesii were the next most common spe- of Calocedrusdecurrens and Abies concolortrees cies, afterPinus ponderosa , respectively.The av- harvestedwas low, averaging13 percentand 33 erageDBH of theQuercus kelloggii was thesmall- percent,respectively (Table 3). estof the species found in themixed conifer stands, The followingcomments were written by George the coniferwith the smallestaverage DBH was Sudworthin the originalfield notebooks and in- Calocedrusdecurrens (Table 1). clude informationabout regeneration and impacts Pinus lambertiana, Pseudotsuga menziesii, and fromearly European settlers (Sudworth 1899). Abies concolorall had similaraverage DBH's of September27, 1899.Near Beech Sawmill(above approximately105 cm whereasPinus ponderosa Placerville)on Big Iowa Canyon.No reproduction and Calocedrusdecurrens had averageDBH's of (manzanitabrush) but abundant a fewyards distant. approximately82 cm. Quercuskelloggii accounted Grazed,no humus,all treesfire marked. foran averageof 6 percentof standstocking. No September28, 1899. South of Blair Sawmill snagswere recorded in the averagemixed conifer (near Sly Park) on summitof ridge.All touched stands(Table 2). withfire, humus 1-2 inchesin spots.Ample repro- Four of the "average" mixedconifer stands in- ductionof all speciesin patches. ventoriedby Sudworthwere in theprocess of being September30, 1899. Sample on big hill south harvestedin 1899. Sudworth'snotebooks recorded west of GrizzlyFlat 0.5 mile. Humus all burned thediameter and speciesof all treesharvested and off. also recordedthe same information on all treesthat October5, 1899.2 mileseast of Whitmore's Mill remainedafter the harvesting operation. All of the Pseudotsugamenziesii trees in these Table 3. AverageAmount Harvested in 4 "average" MixedConifer Stands Located in the Centraland Table 2. Characteristicsof Snags Foundin Mixed NorthernSierra Nevada in 1899. Coniferand Red Fir Stands in the Central and North- ernSierra Nevada in 1899. Basal Basal Trees/ area area DBH of Density Average ha cut cut cut trees Average range Average basal (per- (per- (m2/ cut density(snags/ DBH area Tree cent) cent) ha) (cm) Stand type (snags/ha)ha) (cm) (m2/ha) Abiesconcolor 33.3 48.2 3.9 127 Averagemixed Calocedrusdecurrens 12.5 12.5 1.6 88.9 conifer 0 0 0 0 Pinuslambertiana 87.5 92.8 17.1 104.3 Largemixed Pinusponderosa 57.9 64.2 64.3 103.3 conifer 5 0-10 108.7 4.6 Pseudotsugamenziesii 100.0 100.0 27.2 101.7 Redfir 17.5 0-60 57.3 4.5 Quercuskelloggii 25.0 38.1 2.9 86.4

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Fig.1. El Doradocounty, 1899. Opposite Snyder and Sherman's Ranch. Yellow pine (mixed conifer) forest on south slopeof Silver Fork. Ponderosa pine 91-193 cm (36-76 inches) in diameter,46-50 m high(150-165 feet), clear 8- 11m (25-35feet), ten in 0. 1 ha (0.25acre), 3-5 whitefir {Abies concolor ) same size. Cattle grazed.

(Mill Creek,near Volcano), representingno cut rizes all standcalculations for the large mixed co- stumpage,rolling flat 1000 feetabove creekbot- niferstands. tom. No humus,frequent burning destroyed all. Abies concolorwas the most commonspecies Abundantreproduction of pines and cedar 5-8 comprising46 percentof totalstocking, but only yearsold, mostlyunder 4. accountingfor 34 percentof totalbasal area be- October5, 1899. Near Whitmore'sMill but in cause of theirrelatively small diameters. The larg- shallowravine. Taken as a wholemill operator es- est treeinventoried in thesestands was a Pseudo- timatesoutput 10-20 thousandper acre.Abundant tsugamenziesii and it had a DBH of 188 cm (74 reproductionof all species.Taxus brevi/olia Nutt., inches).Pseudotsuga menziesii made up only 16 dogwood,and Acer macrophyllum Pursh abundant. percentof the trees/habut contributed24 percent Humusin part3-6 inches. of thebasal area ofthe stands because of their large Large mixed coniferstands. The four large size. Abies concolortrees were muchmore com- mixedconifer stands were dominated by largetrees mon in thelarge mixed conifer stands when com- of severalspecies and theaverage quadratic mean pared to the "average" mixedconifer stands (46 diameterat breastheight was 110 cm (43 inches) percentstocking versus 3 percentstocking, respec- forall treesabove 30.5 cm DBH. Averagetree den- tively). sitywas 235 trees/ha(94 trees/acre)(range 160- Abiesconcolor, Abies magnifica , and Calocedrus 300 trees/ha).Average basal areawas large215 m2/ decurrenswere the smallest trees with average qua- ha (923 ft2/acre)(range 188-232 m2/ha). The stands draticmean diameters of approximately93 cm. Pi- inventoriedby Sudworthwere relatively open and nusponderosa and Pinus lambertianawere larger dominatedby largetrees (Fig. 1). Table 4 summa- withaverage diameters of approximately112 cm,

This content downloaded from 169.229.32.36 on Wed, 24 Jun 2015 21:50:04 UTC All use subject to JSTOR Terms and Conditions 2000] STEPHENS:MIXED CONIFER AND RED FIR FORESTSTRUCTURE AND USES IN 1899 47 Table 4. AverageCalculations of GeorgeSudworth's 4 Large MixedConifer Stands in the Centraland NorthernSierra Nevada in 1899(Standard Error). * Averagevalue for all stands. Basalarea Percentof (m2/ha) Trees/ha DBH (cm) totalbasal Percentof Tree [215]* [235]* [110]* area trees/ha Abiesconcolor 72.3 107.5 97.3 34 46 (20.0) (38.2) (6.4) Calocedrusdecurrens 26.7 32.5 92.0 12 14 (9.5) (6.3) (17.4) Pinuslambertiana 33.4 27.5 109.1 16 12 (18.3) (4.8) (20.5) Pinusponderosa 10.5 10.0 115.3 5 4 (10.5) (10.0) (0) Pinusjejfreyi 13.8 10.0 120.4 6 4 (11.4) (7.1) (21.4) Pseudotsugamenziesii 51.8 37.5 123.7 24 16 (45.4) (31.2) (11.8) Abiesmagnifica 6.2 10.0 88.9 3 4 (612) 0O0) (0)

and thelargest trees were Pinus jejfreyi and Pseu- Fork (AmericanRiver) in rich bottombench (at dotsugamenziesii with averagediameters of ap- point where a littlestream enters Silver Fork). proximately122 cm. Quercuskelloggii was notre- Dense firand cedaron outskirts,no seedlingswith- cordedin any of thelarge mixed conifer stands. in. Humus2-3 inches,cattle grazing and sheep. Snag densityaveraged 5/ha with a range0-10/ September21, 1899. 1.5 milessouth of Merzns, ha (Table 2). Averagesnag quadraticmean diam- across(west) of Dark and MultonCanyons (where eterwas 109 cm and snag averagesnag basal area Georgetownroad crosses). South slope of South was 4.6 m2/ha. Fork of the ConsumnesRiver (?) (Sudworthin- The followingcomments were written by George cludedthe? mark and was probablyreferring to the Sudworthin the originalfield notebooks and in- AmericanRiver). Abundant reproduction of all spe- clude informationabout regeneration and impacts cies 1-12 yearsold, all firemarked 15 yearsback. fromearly European settlers (Sudworth 1899). Humus1 .5 inchesdeep, soil sandyloam withrock. September3, 1899. 12-15 mileswest of Bloods, northslope of MokelumneRiver. 30 concolor2-8 Red fir stands.Sudworth sampled four red fir inchesdiameter, 100 under6 inches.5 sugarpine standsduring this inventoryand all standswere under2 feethigh. Thickets of Acer oblusifobium dominatedby Abies magnifica.The averagequa- ( glabruml). draticmean diameterat breastheight was 77 cm September8, 1899. South slope of Bear River, (30 inches)for all treesinventoried. Average tree one halfway up slope.Seedlings of all in spotsnear densitywas 433 trees/ha(173 trees/acre)(range blue ceonothuswhen protected from tramping of 180-610 trees/ha)for trees greaterthat 28 cm cattle.No sheephere, but no humus.Abundant blue DBH. Averagebasal area was 202 m2/ha(867 ft2/ ceonothuschaparral. acre) (range98-286 m2/ha).Table 5 summarizesall September9, 1899. Southlower slope of Silver standcalculations for the red firstands.

Table 5. AverageCalculations of GeorgeSudworth's 4 RedFir Stands in the Central and Northern Sierra Nevadain 1899(Standard Error). * Averagevalue for all stands. Basalarea DBH Percentof (m2/ha) Trees/ha (cm) totalbasal Percentof Tree [202]* [433]* [77]* area trees/ha Pinusjejfreyi 32.2 25.0 128.1 16 6 (32.2) (25.0) (15.1) Abiesmagnifica 136.2 272.5 80.1 67 63 (55.3) (93.5) (10.9) Pinusmontícola 11.3 30.0 66.4 6 7 (8.0) (19.2) (19.4) Pinuscontorta 12.6 70.0 47.8 6 16 (12.6) (70.0) (12.0) Tsugamertensiana 9.3 35.0 58.0 5 8 (9.3) (35.0) (14.5)

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Fig.2. Amadorcounty, 1899. Near sawmill 5-6 km(3-4 miles)below dam on Bear River. Forest fire in fir and pine, killedall seedlings,just started.

The largesttrees in thered fir stands were Abies down on SilverFork (near SilverLake and Kirk- magnificaand Pinus jeffreyi with DBH's of 160 cm wood). Sheep grazing,no reproduction.Scattered (63 inches).Abies magnifica was themost common bunchesof blue ceonothus.Earth bare, rock and treein the standsaccounting for 63 percentof all gravel. treesinventoried and 68 percentof averagestand September7, 1899. Northside of ThimblePeak basal area (Table 5). The nextmost common tree (west of KirkwoodMeadow). On volcanic and foundwas Pinus contortawhich accounted for 16 granite.No humus,grazed by sheep.Dense shade percentof all treesbut only contributed to 6 percent in part,no reproduction.2 Abies magnifica down. of basal area because of thesmallest DBH of any September13, 1899. On Rockyflat between Ly- speciesin thisforest type. ons andBlakley (south fork Silver Creek, west side Snag densityaveraged 17.5 per ha witha range of PyramidPeak). Abundant1 year firseedlings, 0-60 per ha (Table 2). Average snag quadratic 50 firunder 10 feet,20 Murr{Pinus contorta ) pines meandiameter was 57 cm (22 inches)and average 2-10 feet,5 Pimo {Pinusmontícola) 1-3 feet.Hu- snagbasal area was 4.5 m2/ha. mus 1 inch.Cattle grazed, no sheepwithin 5 years. The followingcomments were written by George Sudworthin the originalfield notebooks and in- Discussion clude informationabout regeneration and impacts fromearly European settlers (Sudworth 1899). Sudworth's noted recent evidence of firein many September2, 1899. On foothill(above) Bear standsand believed fireswere ignitedby sheep Meadow,north fork of StanislausRiver. No graz- herdersto increaseforage production and by log- ing,40 youngtrees under 10 inchesdiameter. Hu- gersto consumeslash fuels(Fig. 2). This burning mus 4-6 inchesdeep, no herbaceousgrowth. 75- apparentlydid not spreadextensively because fire 100 seedlings2-10 inches. scaranalysis in theSierra Nevada have documented September7, 1899. On southslope 4-5 miles the almostcomplete removal of surfacefires in

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Fig.3. Calvarascounty, 1899. About 3 km(2 miles)south west of Bloods. Sandy ridge grazed clean and bare by sheep,gullying.

many mixed coniferforests in the 1860-1 870's seedlingswere reportedlyalso trampledby live- (Kilgoreand Taylor1979; Swetnamet al. 1990; stock. Swetnamet al. 1992; Caprio and Swetnam1995) Early loggingoperations had a dramaticeffect at thesame time burning was reportedlybeing used on the species compositionand diameterdistribu- by loggers and sheep herders(Sudworth 1900; tionsof mixedconifer stands sampled by Sudworth McKelvey and Johnston1992; Stephensand El- (Table 3). The majorityof thePinus spp. and Pseu- liott-Fisk1998). dotsuga menziesiiwere harvestedin the stands Regenerationwas notedin themajority of "av- leavinglarge amounts of Calocedrusdecurrens and erage" mixedconifer stands. More site resources Abies concolor(Fig. 4). This typeof loggingop- (light,water, nutrients) were probably available for erationhas been describedas "high-grading"be- regenerationin the"average" mixed conifer stands cause of thepreference for large trees of particular becauseof their lower stocking and basal areas.Re- genera.In thisperiod it was commonfor all mer- generationin mixed coniferforests probably oc- chantabletrees to be removedduring logging op- curredprehistorically when small gaps werecreated erations(Laudenslayer and Darr 1990). Abies con- by theinteraction of fireand locallyhigh fuel loads color and Calocedrusdecurrens were therefore left (Stephenset al. 1999). becausethey were of relatively low economicvalue Regenerationwas noted in half of the red fir late in the 19thcentaury. stands and livestockgrazing was noted in all Earlylogging operations coupled with a national stands.Sudworth noted that in somehigh elevation firesuppression policy that began in theearly 20th sitessheep were actually grazing on coniferseed- centuryfavored shade tolerant species such as Cal- lings(Sudworth 1899). Manyphotos in thecollec- ocedrus decurrensand Abies concolor. Climate tionshow complete bare mineralsoils (Fig. 3) and changesover this period (wetter than average) may

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Fig.4. El Doradocounty, 1899. Forest logged out 5-6 yearsago. Sugar pine (Pinus lambertiana ), large ponderosa pine{Pinus ponderosa ), Douglas-fir (Pseudotsuga menziesii ) taken out, Kellogg oak ( Quercus kelloggii ) and incense cedar( Calocedrusde cur re ns), 12-25 per ha., remain (5-10 per acre). Reproduction ofincense-cedar, ponderosa pine, whitefir, and Douglas-fir abundant 2-9 m high(6-30 feet),2-8 yearsold. Ground grazed. Half mile south of Blairs Millat Sly Park. Humas 4-10 cmdeep (2-4 inches).Soil deep brown, sandy loam. have also led to an increasein treedensities in Si- sitiesin thestands, particularly in thered firforest erraNevada forests. type.Tree density was also muchhigher in thered The managementof Quercuskelloggii is receiv- firforest type when compared to themixed conifer ingincreased attention in theSierra Nevada Frame- forests.Snag basal area was almostidentical in the workProject (SNFP) EnvironmentalImpact State- largemixed conifer stands and red firstands (4.6 mentbecause several rare species such as theCal- m2/haand 4.5 m2/ha,respectively). Snag densities iforniaspotted owl and Pacificfisher use thisspe- foundin thelarge mixed conifer stands are on the cies for foragingand denninghabitat (USDA lowerend of the current requirements for California 2000). Quercuskelloggii is shade intolerant,and spottedowls (Verneret al. 1992). Verylittle snag therefore,would have difficultyliving in areas informationexists for red firforests making it dif- dominatedby largemixed conifers because it can ficultto comparethis historic data to contemporary be over-toppedand killedwhich is one explanation data. of whyit was notrecorded in anylarge mixed co- The averagebasal area recordedin the mixed niferstands. Quercus kelloggii did contributeto 6 coniferstands is high,even for those labeled as percentof average stand stockingon the less "average." The SNFP EnvironmentalImpact Re- stocked"average" mixedconifer stands because portis definingdesired conditions in mixedconifer thesestands were composed by smallertrees, and forestsas havingbasal areas below 70 m2/ha(300 therefore,more site resources were probably avail- ft2/acre).The largemixed conifer stands Sudworth able forthe oaks. inventoriedhad overthree times this basal area and Therewas a greatdeal of variabilityin snagden- the"average" mixed conifer stands had doublethe

This content downloaded from 169.229.32.36 on Wed, 24 Jun 2015 21:50:04 UTC All use subject to JSTOR Terms and Conditions 2000] STEPHENS:MIXED CONIFER AND RED FIR FORESTSTRUCTURE AND USES IN 1899 51 basal area. Some areas of mixedconifer forest in Conclusion theSierra Nevada have the ability to producemuch The mixed coniferstands trees. sampledby George larger Sudworthin 1899 weredominated by largetrees at The mean diameterof the average quadratic relativelylow densities.Shade intolerantspecies, standsfrom this largemixed conifer study(110 cm) Pinus ponderosa, dominatedthe "av- conifer particularly is equal to thoserecorded in the 8 mixed erage" mixed coniferstands whereasthe large standsin thesouthern Sierra Nevada (110 cm) for mixedconifer stands were composed of shadetol- all treesgreater than 30.5 cm DBH. (Stephensand erant,intermediate, and shadeintolerant species. Elliott-Fisk1998). OmittingSequoiadendron gi- Earlyharvesting operations removed the major- ganteum(Lindley) Buchholz (giant sequoia) data ityof the economicallyviable species ( Pinusspp. fromthe fourSequoiadendron giganteum -mixed and Pseudotsugamenziesii) and lefta largeamount coniferstands in the southernSierra Nevada pro- of Calocedrusdecurrens and Abies concolor.This duced an averageDBH of the remainingtrees of practicecoupled with fire suppression policies ini- 111 cm whichis also verysimilar to thoserecorded tiatedat thebeginning of the20th century promoted above. theestablishment and growthof shadetolerant spe- Averagetree density was higherin thesouthern cies. SierraNevada whencompared to the centraland Therewas a largeamount of variabilityin snag northernSierra (278 trees/hacompared to 235 densities,particularly in thered firstands. The red trees/ha,respectively). Average stand basal area firstands had the highesttree densities and Abies was also higherin the mixedconifer stands from magnificadominated in thesestands. the southernSierra Nevada whencompared to the largemixed conifer stands from this study (27 1 m2/ Acknowledgments ha versus215 m2/ha,respectively). Since the av- I posthumouslythank George Sudworth and his inven- erageDBH was equal in themixed conifer stands torycrew for collecting the original data analyzed in this the increasein basal area is a resultof increased paper.I amgrateful toCraig Olsen for introducing meto I wishto stockingin thesouthern Sierra Nevada (18 percent GeorgeSudworth's field notebooks. givespecial recognitiontoNorma Kobzina, Librarian atthe University higher). ofCalifornia Biosciences and Natural Resources Abies concolorwas rarein the Library, very "average" forher assistance on thisproject. mixedconifer stands but was themost common tree of in thelarge mixed conifer stands. Low amounts LiteratureCited Abies concolor in the "average" mixed conifer standsmay have occurredbecause these stands or Bonnicksen,T.M. andE. C. Stone.1982. Reconstruction were probablyless developed(younger) they ofa coniferforest in drierlocations which would have presettlementgiant sequoia-mixed mayhave been communityusing the aggregation approach. Ecology favoredpines over true fir species. In thesouthern 63(4):1134-1 148. Sierra Nevada Abies concolorcontributed to 40 Caprio,A. C. and T. W. Swetnam.1995. Historic fire percentof averagestand stocking and 28 percent regimesalong an elevationalgradient on thewest of averagestand basal area (Stephensand Elliott- slope of the SierraNevada, California, in J. K. onfire in wil- Fisk 1998).In thelarge mixed conifer stands in this Brown,(ed.), Proceedings: Symposium concolorcontributed to 46 of dernessand park management. USD A ForestService study,Abies percent INT-320.Intermoutain Research Sta- and 34 of Gen.Tech. Rep. averagestand stocking percent average tion,Ogden, UT. standbasal area indicatingthat Abies concolor was Gutierrez,R.J., J. Verner, К. S. McKelvey,В. R. Noon, slightlymore common in thecentral and northern G. N. Steger,D. R. Call, W. S. LaHaye,В. В. SierraNevada stands sampled by George Sudworth. Bingham,and J.S. Senser.1992. Habitat relations Pinus lambertianawas muchmore common in ofthe California spotted owl. USD A ForestService thesouthern Sierra Nevada whencompared to the Gen.Tech. Rep. PSW-133. PacificSouthwest Re- centraland northernSierra Nevada of searchStation, Albany, CA. (19 percent Hartes R. J. and H. T. Harvey.1967. The fire 36 ofbasal areaversus 12 percent veldt, stocking, percent ecologyof sequoiaregeneration. Proceedings of the of stocking,16 percentof basal area,respectively). TallTimbers Fire Ecology Conference, Tallehassee, This differencecan be partiallyexplained by the FL. 7:65-77 presenceof Pseudotsugamenziesii in relatively Kilgore,В. M. and D. Taylor.1979. Fire history of a largeamounts (16 percentof stocking,24 percent sequoiamixed conifer forest. Ecology 60(1): 129- of basal area) in thenorthern Sierra Nevada where- 142. Historical as Pseudotsugamenziesii is notnative to thesouth- Laudenslayer,W. F. and H. H. Darr. 1990. ernSierra Nevada. Both menziesiiand effectsof logging on forests of the Cascade and Sierra Pseudotsuga Nevada of California.Transactions of The lambertianaare classifiedas shadeinterme- Ranges Pinus WesternSection of the Wildlife Society 26:12-23. diate(in betweenshade tolerant and shadeintoler- Leopold,S. A.,S. A. Cain,С. А. Соттам,I. N. Gabriel- ant)and therefore, Pseudotsuga menziesii may have son,and T. L. Kimball.1963. Wildlife management occupiedareas thatPinus lambertianacould have in thenational parks. American Forestry 69:32-35, also dominated. 61-63.

This content downloaded from 169.229.32.36 on Wed, 24 Jun 2015 21:50:04 UTC All use subject to JSTOR Terms and Conditions 52 MADROÑO [Vol.47

McKelvey,К. S. and J.D. Johnston.1992. Historical senatedocument #3. Washington D.C. Government perspectiveson forestsof the Sierra Nevada and the printingoffice. TransverseRanges of southern California: Forest con- Swetnam,T. W.,C. H. Baisan,P. M. Brown,A. C. Ca- ditionsat theturn of the century, inJ. Verner et al. prio,and R. Touchan.1990. Late Holocene fire and (eds.),USDA ForestService Gen. Tech. Rep. PSW- climatevariability in giant sequoia groves. Bulletin 133.Pacific Southwest Research Station, Albany, CA. ofthe Ecological Society of America 71(2): 342. Millar,C. I. andW. B. Woolfenden.1999. The role of , Baisan,С. H.,Caprio, A. С., Touchan,R., and climatechange in interpretinghistorical variability. P.M. Brown.1992. Tree ring reconstruction ofgiant EcologicalApplications, vol. 9:1207-1216. sequoiafire regimes. Final report to Sequoia-Kings Parsons,D. J.and S. H. DeBendeetti.1979. Impact of Canyonand YosemiteNational Parks. Cooperative firesuppression on a mixed-coniferforest. Forest agreementDOI 8018-1-002,Laboratory oftree ring Ecologyand Management 2(l):21-33. research,University ofArizona, T'icson, AZ. Rundel,P. W.,D. J.Parsons, and D. T. Gordon.1977. UnitedStates Departmentof Agriculture.2000. In Montaneand subalpine vegetation ofthe Sierra Ne- Press.United States Forest Service Sierra Nevada vadaand Cascade Ranges, in M. F. Barbourand J. FrameworkProject Draft Environmental Impact Maor(eds.), Terrestrial vegetation ofCalifornia, John Statement.Pacific Southwest Region, Mare Island, Wileyand Sons. CA. Stephens,S. L. 1998.Effects of fuelsand silvicultural van Wagtendonk, J.W. 1996.Use of a deterministicfire treatmentson firebehavior in mixed conifer growthmodel to testfuel treatments. Sierra Nevada potential final to vol. forestsof the Sierra Nevada, CA. Forest Ecology and EcosystemProject, report congress, II, Management105:21-34. Assessmentsand ScientificBasis forManagement , D. Dulitz,and R. E. Martin.1999. Giant se- Options.Davis: University ofCalifornia, Centers for in selection ofthe Waterand Wildland Resources. quoiaregeneration group openings Vankat,J. L. andJ. Major. 1978. in southernSierra Nevada. Forest Ecology and Manage- Vegetationchanges ment120:89-95. SequoiaNational Park, California. Journal of Bioge- ography5:377-402. and D. E. Elliott-Fisk.1998. Sequoiadendron K. S. mixedconifer forest structure in 1900- Verner,J., McKelvey,B. R. Noon,R. J.Gutier- giganteum- rez,G. I. Gould,and T. W.Beck. 1992. The Cali- 1901from the southern Sierra Nevada, CA. Madroño, fornia Owl:A technicalassessment ofits cur- vol.45(3):221-230. Spotted rentstatus. USDA ForestService Gen. Tech. Rep. Sudworth,J. B. 1899.Unpublished field note books of PSW-133. Pacific Southwest Research Station, Alba- SierraNevada forest reserve inventory. (University of ny,CA. California,Berkeley, Bioscience and NaturalRe- Weatherspoon,C. P. and C. N. Skinner.1996. Fire sil- sourcesLibrary) viculture relationshipsinSierra forests. Sierra Nevada . 1900.Stanislaus and Lake Tahoe forest reserves, EcosystemProject, final report to congress,vol. II, California,and adjacent territory. In Annual reports Assessmentsand ScientificBasis forManagement ofthe Department ofthe Interior, 21st report of the Options.Davis: University ofCalifornia, Centers for U.S. GeologicalSurvey, 56th Congress, 2nd session, Waterand Wildland Resource.

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