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Appendix 1. Detailed Methods
Aboveground stand characteristics
To quantify carbon (C) and nitrogen (N) pools for individual tree components, we used
component-specific equations developed by Alexander and others (2012) and multiplied the
component biomass by the C and N concentration (foliar C and N concentrations reported in this
study and wood, bark, and twig values obtained from H. Alexander, unpublished). No
component-specific allometry data was available for shrub foliage, so we estimated foliage to be:
total biomass – (branch + stem). For a portion of the smallest shrubs, this calculation yielded a
negative number, possibly due to our shrubs being smaller than those used to develop the
equations. For these shrubs, we calculated component-specific biomass by multiplying the total
biomass by an estimated proportional contribution of foliage and stems (Mack and others 2008).
Shrub C and N concentration data was obtained from H. Alexander (unpublished). Standing dead
tree and shrub biomass was calculated the same way as live biomass, except that the wood
biomass values were multiplied by 0.5 to account for the observation that many dead trees had
snapped, losing the crown and upper stemwood, which we did not quantify in this study. We also
assumed that wood C and N concentration did not differ between live and dead wood and applied
the same values to both.
Tree cross-sections used for aging the stands and estimating aboveground tree net primary
productivity (ANPPtree) were collected at diameter at breast height (DBH, 1.4 m) from a total of
15 trees per species. In the lab, cross-sections were dried and sanded to obtain a smooth, clear
surface, then scanned. Ring number and width were determined using WinDendro software
(Regent Instruments Inc., Quebec, Canada). To calculate ANPPtree, the mean width of the
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preceding 5 years’ growth was used with the wood allometric equations to estimate secondary
growth.
To estimate specific leaf area, a subsample of approximately 100 fresh spruce needles of the
current year’s growth and 10 fresh birch leaves were scanned using WinRhizo software (Regent
Instruments Inc., Quebec, Canada). For the spruce needles, projected leaf area estimates were
multiplied by 1.55 to account for their rhombus shape (Bond-Lamberty and others 2003). For
nutrient analysis, foliage was dried at 60oC, then ground using a wiley mill (Thomas Scientific
model 3383-L10, Swedesboro, NJ). A portion of this material was analyzed for percent C and N
using a Costech Analytical ECS 4010 Elemental Analyzer (Valencia, CA) at the University of
Florida and an additional subsample was shipped to the Louisiana State University AgCenter for
analysis of total phosphorus (P), calcium, magnesium and potassium. Briefly, 5 mL of
concentrated nitric acid was added to 0.5 g of ground sample. After 50 minutes, 3 mL of
hydrogen peroxide was added and the sample was left to digest for 2.75 hours on a heat block.
Samples were then cooled and diluted, then run on a Spectro ARCOS iCAP inductively coupled
plasma spectrometer (Germany).
Downed woody debris was quantified using the line intercept method (Brown 1974) along three,
10 m transects placed at random locations across each plot. Fine woody debris was categorized
into 5 size classes and the number of intercepts of each of class were recorded and converted to
wood mass using multiplier values reported in Nalder and others (1997). The diameter of coarse
woody debris (≥ 7 cm) was also recorded and converted to an area basis as outlined in Ter-
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Mikaelian and others (2008). Biomass estimates were converted to C and N pools using the same
concentration information used to estimate standing tree wood biomass.
Soil Characterization
To estimate moisture content, a weighed subsample of each composite SOL sample was dried at
60oC and each mineral sample at 110oC, then re-weighed. Subsamples of both horizon types
were dried at 60oC and ground prior to analysis of C and N concentration using the same
approach detailed previously for the foliage and litter analysis. Exchangeable base cations were
measured by mixing 50 mL of 1 M ammonium chloride (Robertson and others 1999) with 5 g of
field moist organic horizon sample and 10 g of mineral soil. Samples were shaken for 1 h on a
shaker table, then filtered through a GF/A filter via vacuum filtration and frozen until analysis at
the Louisiana State University AgCenter (Baton Rouge, LA) on a Spectro CIROS inductively
coupled plasma spectrometer (Germany). Mehlich P concentration was determined using a
double acid (0.05 M hydrochloric acid (HCl) + 0.0125 M sulfuric acid) extraction procedure,
with 20 mL of acid solution added to 5 g of air dried soil and shaken for 5 minutes (Kuo 1996).
Samples were then filtered through Whatman No. 5 filter paper and P concentration was
measured colorimetrically using a BioTek Instruments PowerWave XS microplate reder
(Winooski, VT). Soil pH was measured using a Thermo Scientific Orion 2 Star pH meter. For
organic soils, 5 g of air-dried material were mixed with 50 mL of deonized water in a cup and for
mineral soils 20 mL of water were added. Each sample was well mixed, allowed to equilibrate
for 30 minutes, then pH was recorded once the value stabilized.
To calculate potential N mineralization and nitrification, ammonium (NH4+) and nitrate (NO3
-)
were extracted from a subsample of the initial composite soils, as well as for the 30-day and 90-
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day incubations detailed in the Soil C and N fluxes section of the main text and below. Briefly, 5
g of moist organic soil and 10 g of mineral soil were mixed with 50 mL of 2 M potassium
chloride for 1 hr (Robertson and others 1999). Samples were then filtered through a GF/A filter
via vacuum filtration and immediately frozen until analysis on an Astoria-Pacific International
Autoanalyzer (Clackamas, OR) at the University of Florida.
Near total element digestion was performed on all mineral soils (0-10 cm plot composite samples
and all deeper, ≤ 1 m mineral soil increments) by ALS Minerals (Reno, NV) using a four acid
near total digestion method. First, 0.25 g of ground sample was digested with HCl, perchloric,
nitric, and hydrofluoric acids. The residue was brought to volume with dilute HCl, then analyzed
on an inductively coupled plasma atomic emission spectrometer.
To radiocarbon date the soils, green moss was first removed from the SOL surface of black
spruce SOLs using scissors. The core was then cut in half vertically. One half of the core was
carefully cut into 5 cm depth increments and each segment was analyzed for moisture content,
bulk density, and C and N using the same methods as described for SOL processing previously.
The second half was carefully divided into 1 cm depth increments. A small portion of these
segments were then smeared on a piece of paper to qualitatively determine the presence/absence
of charcoal within the increment (Jones and others 2013). When the charcoal layer was found,
the increment containing the charcoal and at least 2 increments on both sides of this increment
were chosen for further processing and radiocarbon analysis. Moss macrofossils were carefully
removed with a dissecting scope from the chosen increments in the black spruce SOLs and a
bulk soil sample (excluding roots) was also obtained. The birch SOL did not contain any
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charcoal or mosses, so only a root-free bulk soil sample was analyzed. Cellulose was extracted
from the macrofossil samples (Gaudinski and others 2005) and all macrofossil and bulk soil
samples were converted to graphite at 650oC with an iron catalyst in a hydrogen atmosphere
(Vogel and others 1987). Graphite samples were then sent to the UC Irvine W.M. Keck Carbon
Cycle Accelerator Mass Spectrometry Laboratory for analysis. Values obtained for the bulk and
macrofossil samples from a given depth increment in spruce stands differed little and here we
only report the bulk values. Also, we were unable to keep the black spruce SOL core from block
C intact and therefore it was excluded from the analysis.
Ion exchange resins were rinsed with deionized water after returning to the lab and refrigerated
until analysis. In preparation for extraction, resin bags were rinsed for 20 seconds with nano-pure
water to remove all remaining soil particles. Each bag was placed in a tube with 30 mL of a
mixture of 0.1 M HCl and 2 M sodium chloride and shaken for 1 hr on a shaker table. Extractant
was then drip-filtered through a Whatman GF/A filter and frozen until further analysis.
Phosphate was measured colorimetrically within 1 week of extraction using a spectrophotometer
microplate reader (PowerWave XS Microplate Reader, Bio-Tek Instruments, Inc., Winooski,
VT) using the ascorbic acid molybdenum-blue method (Murphy and Riley 1962). The remaining
sample was frozen until analysis of NH4+ and NO3
- using a segmented flow autoanalyzer
(Astoria-Pacific, Inc., Clackamas, OR) at the University of Florida.
Soil C and N fluxes
To estimate soil carbon dioxide (CO2) respiration (90-day) and potential net N mineralization
and nitrification (30- and 90-day), an approximately 3 cm x 3 cm intact piece of each SOL
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horizon was removed from each intact core so as to include the entire vertical length of the
horizon. Subsamples were then placed in a 32 oz. mason jar with glass beads underlying
perforated aluminum foil on the bottom of the jar. For each horizon, the subsamples originating
from the 3 field sampling locations within a given plot were put into a single jar, creating 1
composite sample per plot, per horizon. For each 10 cm mineral soil core, a small portion of soil
was removed using a spatula vertically along the length of the core. Once in jars, deionized water
was added to each sample to approximate field capacity and samples were placed in the dark at
15oC. Soil CO2 respiration was measured using an automated soil incubation system equipped
with an infrared gas analyzer (Li-820, Licor, Inc, Lincoln, NE) (Bracho and others unpublished).
To estimate 14CO2, CO2 was first removed from each jar by scrubbing with magnesium
perchlorate and soda lime for 5 min. Jars were then left at 15oC for 3-5 days to allow CO2 to
accumulate. The headspace air was then pulled through a zeolite molecular sieve trap (Alltech
13X, Alltech Associates, Deerfield, IL) to absorb the CO2, baked, reduced to graphite (Hicks
Pries and others 2013), then sent to the UC Irvine W.M. Keck Carbon Cycle Accelerator Mass
Spectrometry Laboratory for final analysis.
REFERENCES
Alexander HD, Mack MC, Goetz S, Beck PSA, Belshe EF. 2012. Implications of increased
deciduous cover on stand structure and aboveground carbon pools of Alaskan boreal forests.
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Bond-Lamberty B, Wang C, Gower ST. 2003. The use of multiple measurement techniques to
refine estimates of conifer needle geometry. Canadian Journal of Forest Research-Revue
Canadienne De Recherche Forestiere 33: 101-105.
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Brown JK. 1974. Handbook for inventorying downed woody material. Ogden, Utah, USA:
USDA Forest Service, Intermountain Forest and Range Experiment Station.
Gaudinski JB, Dawson TE, Quideau S, Schuur EAG, Roden JS, Trumbore SE, Sandquist DR,
Oh SW, Wasylishen RE. 2005. Comparative analysis of cellulose preparation techniques for
use with 13C, 14C, and 18O isotopic measurements. Analytical Chemistry 77: 7212-7224.
Hicks Pries CE, Schuur EAG, Crummer KG. 2013. Thawing permafrost increases old soil and
autotrophic respiration in tundra: Partitioning ecosystem respiration using δ13C and Δ14C.
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DA. 2013. Identification of unrecognized tundra fire events on the north slope of Alaska.
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chemical methods. Madison: Soil Science Society of America, p893-894.
Mack MC, Treseder KK, Manies KL, Harden JW, Schuur EAG, Vogel JG, Randerson JT,
Chapin FS, III. 2008. Recovery of aboveground plant biomass and productivity after fire in
mesic and dry black spruce forests of interior alaska. Ecosystems 11: 209-225.
Murphy J, Riley JP. 1962. A modified single solution method for determination of phosphate in
natural waters. Analytica Chimica Acta 26: 31-&.
Nalder IA, Wein RW, Alexander ME, deGroot WJ. 1997. Physical properties of dead and
downed round-wood fuels in the boreal forests of Alberta and Northwest Territories.
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Robertson GP, Coleman DC, Bledsoe CS, Sollins P editors. 1999. Standard soil methods for
long-term ecological research. New York: Oxford University Press.
Ter-Mikaelian MT, Colombo SJ, Chen JX. 2008. Amount of downed woody debris and its
prediction using stand characteristics in boreal and mixedwood forests of Ontario, Canada.
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graphite for AMS. Nuclear Instruments & Methods in Physics Research Section B-Beam
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Appendix 2. Forest Stand Characteristics for Each Spatially Interspersed Block of Black Spruce and Paper Birch across the Study Site
Black spruce Paper birch
Block A Block B Block C Block A Block B Block C
Tree age (years) 45.4(0.7) 38.5(2.3) 44.0(1.0) 49.8(0.8) 47.9(0.8) 46.8(1.3)Density (stems m-2) All stems 8.22(1.43) 8.29(0.94) 5.43(0.47) 0.56(0.05) 1.21(0.13) 1.04(0.08) Plot target species BD 4.72(13.2) 4.68(1.04) 1.7(0.32) 0.02(0.02) 0.06(0.04) 0.1(0.01) DBH 2.95(0.44) 3.05(0.32) 3.14(0.41) 0.50(0.05) 0.88(0.09) 0.80(0.10) Other species BD 0.03(0.01) 0.04(0.01) 0.002(0.002) 0.02(0.01) 0.23(0.13) 0.11(0.02) DBH 0.22(0.05) 0.25(0.11) 0.06(0.01) 0.01(0.004) 0.02(0.01) 0.02(0.01) Tall shrubs 0.31(0.07) 0.27(0.09) 0.58(0.07) 0.00(0.00) 0.02(0.02) 0.10(0.03)Basal area (m2 ha-1) All stems 20.99(3.24) 19.81(1.18) 24.60(2.85) 29.41(1.58) 25.67(1.62) 30.82(4.27) Plot target species BD 4.10(1.15) 4.07(0.90) 1.41(0.20) 0.005(0.004) 0.03(0.03) 0.02(0.01) DBH 12.84(3.11) 10.98(1.03) 18.04(2.37) 29.31(1.59) 24.43(1.37) 28.76(4.28) Other species BD 0.18(0.09) 0.12(0.05) 0.002(0.002) 0.07(0.06) 0.20(0.09) 0.10(0.03) DBH 2.42(0.69) 2.38(1.16) 1.39(0.49) 0.02(0.01) 0.12(0.07) 0.08(0.05) Tall shrubs 1.44(0.48) 2.26(.068) 3.76(0.59) 0.00(0.00) 0.89(0.55) 1.85(0.45)Live tree biomass (kg m-2) All stems 4.92(1.08) 4.62(0.54) 6.39(0.91) 12.06(0.80) 9.88(0.67) 12.04(1.81) Plot target species BD 0.22(0.06) 0.21(0.05) 0.08(0.01) 0.0001(0.0001) 0.002(0.002) 0.001(0.001) DBH 3.66(0.83) 3.20(0.29) 5.02(0.64) 12.04(0.80) 9.43(0.59) 11.51(1.79)
Other species BD 0.02(0.01) 0.01(0.004) 0.0001(0.0001) 0.01(0.01) 0.01(0.004) 0.01(0.002) DBH 0.78(0.24) 0.77(0.38) 0.50(0.20) 0.01(0.004) 0.03(0.02) 0.02(0.01) Tall shrubs 0.24(0.10) 0.42(0.14) 0.80(0.20) 0.00(0.00) 0.41(0.27) 0.50)(0.10)
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Appendix 2. Continued
Proportion biomass Plot target species 0.80(0.03) 0.76(0.05) 0.80(0.04) 1.00(0.001) 0.96(0.03) 0.95(0.01) Other species 0.15(0.02) 0.14(0.06) 0.07(0.02) 0.001(0.001) 0.004(0.002) 0.002(0.001) Tall shrubs 0.04(0.02) 0.10(0.03) 0.12(0.02) 0.00(0.00) 0.04(0.03) 0.05(0.01)
All values are mean (SE), n = 5 plots per block.
Trees ≥ 1.4 m in height are included in the rows labeled DBH (diameter at breast height) and trees < 1.4 m tall are included in the BD
(basal diameter) category.
Other species includes all measure trees that were not the focus species studied within the given plot.
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Appendix 3. Total Elemental Concentrations for Mineral Soils (0-100 cm) for Studied Black Spruce and Alaska Paper Birch Stands
Black spruce
Soil Depth Increment (cm)0-10 10-20 20-30 30-40 40-50 50-60 60-70 70-80 80-90 90-100
C % 2.80(0.34) 2.12(1.09) 1.20(0.43) 1.07(0.42) 1.63(0.84) 1.57(1.03) 0.64 ND ND NDN % 0.16(0.02) 0.13(0.06) 0.07(0.02) 0.07(0.02) 0.10(0.05) 0.09(0.05) 0.05 ND ND NDAg (ppm) 12.8(3.6) 11.1(4.5) 19.4(3.2) 27.6(3.5) 19.0(7.5) 17.5(1.6) 17.3 ND ND NDAl % 6.66(0.65) 6.49(0.49) 6.50(0.46) 6.73(0.48) 6.90(0.44) 6.88(0.35) 6.54 ND ND NDAs (ppm) 11(1) 16(4) 13(1) 13(2) 13(0) 15(1) 16 ND ND NDBa (ppm) 757(64) 752(36) 749(35) 761(38) 807(48) 777(31) 834 ND ND NDBe (ppm) 1.3(0.1) 1.3(0.1) 1.3(0.1) 1.4(0.1) 1.5(0.1) 1.4(0.1) 1.3 ND `ND NDBi (ppm) 1(1) 1(1) 1(1) 1(1) 1(1) 2(1) 2 ND ND NDCa % 1.39(0.32) 1.29(0.20) 1.24(0.19) 1.24(0.18) 1.21(0.12) 1.20(0.15) 1 ND ND NDCd (ppm) 0(0) 1(0) 3(0) 4(1) 2(1) 1(0) 3 ND ND NDCr (ppm) 86(10) 82(6) 77(7) 79(3) 82(4) 80(3) 74 ND ND NDCu (ppm) 24(6) 31(4) 32(6) 37(2) 38(2) 36(4) 36 ND ND NDFe % 3.46(0.32) 3.78(0.14) 3.61(0.20) 3.73(0.20) 3.84(0.16) 3.89(0.21) 3.66 ND ND NDGa (ppm) 13(3) 17(3) 17(3) 16(3) 20(0) 20(0) 16 ND ND NDK % 1.47(0.14) 1.46(0.14) 1.52(0.14) 1.61(0.21) 1.62(0.16) 1.65(0.18) 1.46 ND ND NDLa (ppm) 33(3) 33(3) 30(6) 30(6) 33(3) 35(5) 30 ND ND NDMg (%) 1.03(0.11) 1.02(0.07) 1.01(0.09) 1.06(0.09) 1.08(0.08) 1.08(0.06) 1.04 ND ND NDMn (ppm) 487(23) 512(33) 533(61) 533(62) 578(26) 563(47) 509 ND ND NDNa % 1.26(0.16) 1.17(0.06) 1.16(0.13) 1.14(0.13) 1.12(0.04) 1.12(0.14) 1.31 ND ND NDNi (ppm) 29(4) 31(3) 30(3) 32(2) 36(3) 35(3) 32 ND ND NDP (ppm) 665(31) 698(117) 624(53) 610(50) 620(26) 673(3) 694 ND ND NDPb (ppm) 15(1) 13(1) 31(12) 18(2) 15(1) 15(0) 14 ND ND NDS % 0.02(0.003) 0.02(0.01) 0.01(0.01) 0.01(0.01) 0.02(0.01) 0.01(0.00) 0.01 ND ND NDSc (ppm) 14(2) 14(1) 13(1) 14(1) 15(1) 15(1) 14 ND ND NDSr (ppm) 188(30) 177(18) 176(22) 176(19) 175(11) 177(12) 201 ND ND NDTi % 0.43(0.04) 0.42(0.03) 0.40(0.03) 0.43(0.03) 0.43(0.03) 0.44(0.01) 0.41 ND ND ND
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V (ppm) 109(12) 109(8) 104(8) 108(5) 113(6) 110(1) 109 ND ND NDZn (ppm) 76(3) 78(2) 84(2) 88(5) 86(7) 80(2) 89 ND ND ND
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Appendix 3. Continued
Paper birchSoil Depth Increment (cm)
0-10 10-20 20-30 30-40 40-50 50-60 60-70 70-80 80-90 90-100C % 4.13(0.76) 1.54(0.32) 0.91(0.27) 0.60(0.12) 0.54(0.13) 0.65(0.14) 0.50(0.04) 0.51 0.59 0.61N % 0.20(0.03) 0.09(0.01) 0.05(0.01) 0.04(0.01) 0.05(0.00) 0.05(0.00) 0.04(0.01) 0.05 0.05 0.06Ag (ppm) 0.5(0.5) 0.70(0.70) 1.3(0.8) 1.4(1.1) 8.4(8.1) 5.6(5.6) 8.9(8.9) 0 0.5 1.7Al % 6.20(0.18) 6.49(0.20) 6.90(0.33) 6.57(0.13) 6.46(0.12) 6.35(0.17) 6.69(0.24) 6.44 6.25 6.7As (ppm) 10(1) 13(1) 13(1) 15(2) 9(5) 10(2) 12(2) 13 19 15Ba (ppm) 677(18) 707(39) 753(52) 713(73) 717(66) 627(107) 680(170) 870 880 930Be (ppm) 1.1(0.0) 1.2(0.1) 1.2(0.1) 1.2(0.1) 1.2(0.1) 1.1(0.1) 1.2(0.1) 1.3 1.3 1.3Bi (ppm) 1(1) 1(1) 0(0) 1(1) 1(1) 1(1) 4(2) 0 4 0Ca % 1.44(0.36) 1.36(0.33) 1.50(0.35) 1.53(0.32) 1.34(0.30) 1.51(0.15) 1.61(0.21) 1.72 1.65 1.73Cd (ppm) 0(0) 0(0) 0(0) 0(0) 0(0) 0(0) 0(0) 0 0 0Cr (ppm) 105(27) 98(22) 104(18) 98(12) 86(9) 90(5) 97(14) 78 79 83Cu (ppm) 28(8) 29(6) 33(6) 36(4) 36(5) 51(10) 57(22) 34 33 36Fe % 3.42(0.33) 3.71(0.17) 3.95(0.22) 4.03(0.19) 3.98(0.38) 4.58(0.44) 4.70(0.96) 3.58 3.63 3.74Ga (ppm) 13(3) 20(0) 17(3) 20(0) 20(0) 20(0) 20(0) 20 10 20K % 1.22(0.03) 1.26(0.03) 1.32(0.04) 1.21(0.11) 1.33(0.13) 1.12(0.13) 1.24(0.19) 1.43 1.40 1.48La (ppm) 23(3) 23(3) 33(3) 30(6) 30(0) 27(3) 25(5) 30 30 30Mg (%) 1.01(0.16) 1.06(0.14) 1.18(0.19) 1.16(0.13) 1.08(0.14) 1.23(0.06) 1.29(0.13) 1.13 1.09 1.16Mn (ppm) 459(69) 428(38) 504(59) 540(65) 553(68) 702(105) 789(40) 683 785 778Na % 1.20(0.15) 1.21(0.17) 1.33(0.21) 1.29(0.16) 1.17(0.17) 1.17(0.17) 1.24(0.25) 1.45 1.37 1.47Ni (ppm) 34(10) 36(7) 41(9) 40(9) 38(7) 41(3) 45(6) 37 39 41P (ppm) 593(133) 467(68) 520(136) 507(113) 533(146) 520(150) 620(190) 810 800 780Pb (ppm) 12(1) 64(50) 14(1) 14(2) 23(9) 19(8) 13(1) 15 17 17S % 0.01(0.003) 0.01(0.003) 0.01(0.003) 0.003(0.003) 0.003(0.00) 0.003(0.003) 0.01(0.01) 0.01 0.02 0.01Sc (ppm) 14(1) 14(1) 16(2) 16(1) 15(1) 17(1) 18(3) 14 14 15Sr (ppm) 179(36) 177(33) 197(41) 192(35) 173(32) 169(34) 183(48) 226 216 229Ti % 0.45(0.00) 0.41(0.02) 0.46(0.02) 0.47(0.03) 0.45(0.06) 0.49(0.05) 0.50(0.08) 0.42 0.41 0.42V (ppm) 116(9) 113(12) 125(12) 128(9) 123(14) 144(15) 151(31) 115 114 119Zn (ppm) 68(9) 70(6) 77(6) 75(6) 76(5) 77(4) 82(3) 83 85 92
Mean (SE), n = 3 samples per species, per depth increment. Cells containing ND indicate no data available.181182