uid=HFR,o=lter,dc=ecoinformatics,dc=org all public read doi:10.6073/pasta/45bf2e8b1371c0347edb742d0e7ef684 Shannon Pelini Angus Chen Justine Kaseman Thomas Crowther Audrey Maran https://orcid.org/0000-0003-3640-2549 Researcher 2023 English

Forest floor food webs play pivotal roles in carbon cycling, but they are rarely considered in models of carbon fluxes, including soil carbon dioxide emissions (respiration), under climatic warming. The indirect effects of invertebrates on heterotrophic respiration through interactions with microbial communities are significant and will be altered by warming. However, the interactive effects of invertebrates and warming on microbes and heterotrophic respiration in the field are poorly understood. In this study we combined field and common garden laboratory approaches to examine relationships between warming, forest floor food web structure, and heterotrophic respiration. We found that soil animals can overwhelm the effects of warming (to 5 degrees Celsius above ambient) on heterotrophic respiration. In particular, the presence of higher trophic levels and burrowing detritivores strongly determined heterotrophic respiration rates in temperate forest soils, dictating the ecosystem response to warming. These effects were, however, context-dependent, with greater effects in a lower-latitude site. Without isolating and including the significant impact of invertebrates, climate models will be incomplete, hindering well-informed policy decisions.

arthropods carbon climate change community structure food webs insects soil respiration trophic structure LTER controlled vocabulary organic matter LTER core area Harvard Forest HFR LTER USA HFR default

This dataset is released to the public under Creative Commons CC0 1.0 (No Rights Reserved). Please keep the dataset creators informed of any plans to use the dataset. Consultation with the original investigators is strongly encouraged. Publications and data products that make use of the dataset should include proper acknowledgement.

Creative Commons Zero v1.0 Universal https://spdx.org/licenses/CC0-1.0.html CC0-1.0 https://harvardforest.fas.harvard.edu/exist/apps/datasets/showData.html?id=hf276 Prospect Hill Tract (Harvard Forest), Duke Forest (NC). Coordinates based on WGS84 datum. -79.98 -72.19 +43.53 +35.87 130 340 meter 2013 2013 complete Information Manager Harvard Forest

324 North Main Street Petersham MA 01366 USA

(978) 724-3302 hf-im@lists.fas.harvard.edu Harvard Forest

324 North Main Street Petersham MA 01366 USA

(978) 724-3302 (978) 724-3595 https://harvardforest.fas.harvard.edu

Soil cores (~20cm diameter, organic layer) were collected from warm ant chambers at Duke Forest on 10 July 2013 and shipped to Harvard Forest, where macroinvertebrates (>1cm) were separated, recorded, and added to mesocosms (20 cm tall x 10 cm diameter PVC columns with rubber bottom cap and covered with black landscape cloth secured with rubber band). Mesocosms contained homogenized soils, with macroinvertebrates removed, collected near Harvard Forest warming chambers. Mesocosms were placed in Shaler basement wood shop. Experiment ran 17 July-19 August 2013. We measured leaf decomposition (round leaf litter bags course enough for invertebrate access on the topside; leaves were plucked and air dried from one Harvard Forest maple tree). Mesocosms were harvested at the end of the experiment and all living and dead invertebrate bodies were recorded. Respiration measurements were taken on a LI-6400 (LICOR, Lincoln, NE). Invertebrates were identified broadly into the following categories: spider, centipede, beetle larva morph 1, beetle larva morph 2, millipede, fly larva, caterpillar, earthworm, ant, leaf beetle, beetle. NA's represent missing or not relevant data.

Same as “Duke Forest Common Garden Experiment—Invertebrate”, but soil cores (~20cm diameter, organic layer) were collected from warm ant chambers at Harvard Forest on 1 June 2013. Invertebrates were not included in any mesocosms.

Soil cores (~20cm diameter, organic layer) were collected from warm ant chambers at Harvard Forest on 1 June 2013. Macroinvertebrates (>1cm) were separated, recorded, and added to mesocosms (20 cm tall x 10 cm diameter PVC columns with rubber bottom cap and covered with black landscape cloth secured with rubber band). Mesocosms contained homogenized soils left over from soil cores upon macroinvertebrate extraction. Mesocosms were placed in Torrey greenhouse. Experiment was moved on 25 June 2013 into Shaler greenhouse, where temperatures spiked, causing nearly 100 % mortality, thus ending the experiment. We measured leaf decomposition (round leaf litter bags course enough for invertebrate access on the topside; decomposing maple and birch leaf mix was collected near the warming chambers. Measurements as described in “Duke Forest Common Garden Experiment—Invertebrate.” Invertebrates were identified broadly into the following categories: beetle larva, beetle small ambrosia-like, beetle-rove, beetle-other, tiger beetle morph1, tiger beetle morph2, beetle black, beetle firefly, beetle-Japanese, beetle-weevil, spider-large, spider-medium, spider-small, spider-jump, hemiptera,ant- myrmica, ant-formica, ant- aphaenogaster, ant-tapinonma sessile, ant- temnothorax, ant-lasius, ant-camponotus, opilones (harvestman), homoptera (aphid), millipede, centipede, slug, caterpillar. NA's represent missing or not relevant data.

We filled mesocosms (19 L buckets, 30 cm diameter, 44 cm height) to 10 cm height with topsoil (O and A horizons, Canton loam) that we collected ~100-200 yards away from the chambers, from which we removed roots and macrofauna (e.g., beetles, ants, spiders, millipedes) via sieving (1mm mesh), and homogenized across all mesocosms. We had 5 treatments: 1: no fauna present; 2: only micro/mesofauna present; 3: only macroinvertebrates present; 4: micro/mesofauna and macroinvertebrates present; and 5: micro/mesofauna, macroinvertebrates, and a vertebrate predator present. We removed micro- (e.g., fungi, bacteria) and meso- (e.g., mites, springtails) fauna (Treatments 1 and 3) by autoclaving soil described above at 120°C for 30 minutes. We used red back salamanders (Plethodon cinereu) as the top predator in this experiment (Treatment 4). For treatments that included macroinvertebrates (Treatments 3 and 4), we used ~1.5 g each of live earthworms (Megadrilacea), a common invertebrate engineer and food source for salamanders, and mealworms (Tenebrionidae). Mesocosms were placed in the warm ants warming chambers (1.5-5.5 degrees C above ambient) at Harvard Forest Prospect Hill Tract. Into each mesocosm, we also installed a 10 cm diameter, 4.5 cm high, PVC collar, inserted approximately 1 cm into the soil. Within these collars we collected weekly soil efflux (RH) measurements using a LiCor 6400 (Lincoln, Nebraska). We counted and weighed all surviving macroinvertebrates at the end of the experiment. Mesocosms were present in the chambers for 6 weeks during June-August 2013. Within a few days of being placed in the field, micro/mesofauna removal mesocosms (Treatments 1 and 3) had dense surface fungal colonization while those with fauna (Treatments 2, 4 and 5) did not. We visually estimated the amount of fungal coverage in each mesocosm using a qualitative scale from 0 (no visible fungus) to 10 (no bare soil visible).

Harvard Forest

324 North Main Street Petersham MA 01366 USA

(978) 724-3302 (978) 724-3595 https://harvardforest.fas.harvard.edu https://ror.org/059cpzx98 pointOfContact The Harvard Forest Long-Term Ecological Research (LTER) program examines ecological dynamics in the New England region resulting from natural disturbances, environmental change, and human impacts. National Science Foundation LTER grants: DEB-8811764, DEB-9411975, DEB-0080592, DEB-0620443, DEB-1237491, DEB-1832210. hf276-01-DF-invertebrate.csv Duke Forest common garden experiment -invertebrate hf276-01-DF-invertebrate.csv 19243 811b98ddd166a45e1655b000b1930dd8 1 \r\n column , https://harvardforest.fas.harvard.edu/data/p27/hf276/hf276-01-DF-invertebrate.csv date date of CO2 efflux measurement YYYY-MM-DD 1 day NA missing value week week of the experiment week of the experiment NA missing value tag unique identifier number on metal tag attached to each decomposition bag unique identifier number on metal tag attached to each decomposition bag NA missing value efflux soil efflux (μmol CO2 mol-1) measured using LI-6400 dimensionless 0.000000001 real NA missing value temp mean air temperature delta (difference between chamber and average of 3 ambient reference stations) of Duke Forest warming chambers between April 2010 and July 2013 from which soil cores/invertebrates were extracted celsius 0.00001 real NA missing value chamber DF ant warming chamber from which the invertebrates originated DF ant warming chamber from which the invertebrates originated NA missing value subsample A-D, from 4 opposite sides of each chamber A-D, from 4 opposite sides of each chamber NA missing value mesocosm identifier derived from chamber and subsample identifier derived from chamber and subsample NA missing value core.depth depth of extracted soil core centimeter 0.1 real NA missing value leaf.decomp percent leaf mass lost from decomposition bag dimensionless 0.000000001 real NA missing value invert.biomass.i total mass of macroinvertebrates extracted from soil core and placed in mesocosm gram 0.01 real NA missing value invert.abundance.i total number of macroinvertebrate individuals extracted from soil core and placed in mesocosm number 1 whole NA missing value invert.richness.i total richness, at a broad morphological/order scale, of macroinvertebrates extracted from soil core and placed in mesocosm number 1 whole NA missing value invert.biomass.f total mass of macroinvertebrates that survived to the end of the mesocosm experiment gram 0.001 real NA missing value invert.abundance.f total number of macroinvertebrate individuals that survived to the end of the mesocosm experiment number 1 whole NA missing value invert.richness.f total richness, at a broad morphological/order scale, of macroinvertebrates that survived to the end of the mesocosm experiment number 1 whole NA missing value microbial.biomass.c microbial biomass (organic) carbon, using chloroform fumigation method in the lab of M Weintraub. TOC ug-C/g dry, calculated as difference between fumigated and K2SO4 extracted sample dimensionless 0.0001 real NA missing value microbial.biomass.n microbial biomass nitrogen, using chloroform fumigation method in the lab of M Weintraub. TN ug-N/g dry, calculated as difference between fumigated and K2SO4 extracted sample. dimensionless 0.0001 real NA missing value spider number of spiders extracted from soil core and placed in mesocosm number 1 whole NA missing value centipede number of centipedes extracted from soil core and placed in mesocosm number 1 whole NA missing value millipede number of millipedes extracted from soil core and placed in mesocosm number 1 whole NA missing value maggot number of diptera larva extracted from soil core and placed in mesocosm number 1 whole NA missing value caterpillar number of caterpillars extracted from soil core and placed in mesocosm number 1 whole NA missing value earthworm number of earthworms extracted from soil core and placed in mesocosm number 1 whole NA missing value ant number of ants extracted from soil core and placed in mesocosm number 1 whole NA missing value beetle number of beetles extracted from soil core and placed in mesocosm number 1 whole NA missing value 176 hf276-02-HF-invertebrate.csv Harvard Forest common garden experiment - invertebrate hf276-02-HF-invertebrate.csv 12476 2738894b788c529497286599f39a6e24 1 \r\n column , https://harvardforest.fas.harvard.edu/data/p27/hf276/hf276-02-HF-invertebrate.csv date date of CO2 efflux measurement LI-6400 YYYY-MM-DD 1 day NA missing value week week of the experiment week of the experiment NA missing value tag unique identifier number on metal tag attached to each decomposition bag unique identifier number on metal tag attached to each decomposition bag NA missing value efflux soil efflux (μmol CO2 mol-1) dimensionless 0.000000001 real NA missing value temp mean air temperature delta (difference between chamber and average of 3 ambient reference stations) of Harvard Forest warming chambers between April 2010 and July 2013 from which soil cores/invertebrates were extracted celsius 0.000001 real NA missing value chamber ant warming chamber invertebrates originated from at HF ant warming chamber invertebrates originated from at HF NA missing value subsample A-D, from 4 opposite sides of each chamber A-D, from 4 opposite sides of each chamber NA missing value mesocosm identifier derived from chamber and subsample identifier derived from chamber and subsample NA missing value core.depth1 depth of extracted soil core centimeter 0.1 real NA missing value core.depth2 depth of second extracted soil core taken due to error in collection of first core. If core.depth2 is provided, this was the depth of the core used for the experiment, otherwise core.depth1 was used. centimeter 0.1 real NA missing value leaf.decomp percent leaf mass lost from decomposition bag dimensionless 0.000000001 real NA missing value invert.biomass.i total mass of macroinvertebrates extracted from soil core and placed in mesocosm gram 0.001 real NA missing value invert.abundance.i total number of macroinvertebrate individuals extracted from soil core and placed in mesocosm number 1 whole NA missing value invert.richness.i total richness, at a broad morphological/order scale, of macroinvertebrates extracted from soil core and placed in mesocosm number 1 whole NA missing value beetle number of beetles extracted from soil core and placed in mesocosm number 1 whole NA missing value spider number of spiders extracted from soil core and placed in mesocosm number 1 whole NA missing value hemiptera number of hemipterans extracted from soil core and placed in mesocosm number 1 whole NA missing value ant number of ants extracted from soil core and placed in mesocosm number 1 whole NA missing value opilones number of opilones extracted from soil core and placed in mesocosm number 1 whole NA missing value homoptera number of homopterans extracted from soil core and placed in mesocosm number 1 whole NA missing value millipede number of millipedes extracted from soil core and placed in mesocosm number 1 whole NA missing value centipede number of centipedes extracted from soil core and placed in mesocosm number 1 whole NA missing value slug number of slugs extracted from soil core and placed in mesocosm number 1 whole NA missing value caterpillar number of caterpillars extracted from soil core and placed in mesocosm number 1 whole NA missing value 132 hf276-03-HF-microbe.csv Harvard Forest common garden experiment - microbe hf276-03-HF-microbe.csv 15853 f64a1de399b5939fe8c9cde26ca9430e 1 \r\n column , https://harvardforest.fas.harvard.edu/data/p27/hf276/hf276-03-HF-microbe.csv date date of CO2 efflux measurement YYYY-MM-DD 1 day NA missing value week week of the experiment week of the experiment NA missing value temp temperature of the soil at approximately 8 cm deep celsius 0.000001 real NA missing value tag unique identifier number on metal tag attached to each decomposition bag unique identifier number on metal tag attached to each decomposition bag NA missing value chamber HF ant warming chamber from which the invertebrates originated ant warming chamber invertebrates originated from at HF NA missing value subsample A-D, from 4 opposite sides of each chamber A-D, from 4 opposite sides of each chamber NA missing value mesocosm identifier derived from chamber and subsample identifier derived from chamber and subsample NA missing value core.depth depth of extracted soil core centimeter 0.1 real NA missing value leaf.decomp percent leaf mass change in decomposition bag. Negative values indicate leaf mass was gained. dimensionless 0.000000001 real NA missing value leaf.gain.loss indicates if leaf mass was lost or gained loss leaf mass lost gain leaf mass gained NA missing value microbial.biomass.c microbial biomass (organic) Carbon, using chloroform fumigation method in the lab of M Weintraub. TOC ug-C/g dry soil, calculated as difference between fumigated and K2SO4 extracted sample. dimensionless 0.0000001 real NA missing value microbial.biomass.n microbial biomass nitrogen, using chloroform fumigation method in the lab of M Weintraub. TN ug-N/g dry soil, calculated as difference between fumigated and K2SO4 extracted sample. dimensionless 0.0000001 real NA missing value efflux soil efflux (μmol CO2 mol-1) measured using LI-6400 dimensionless 0.000000001 real NA missing value 177 hf276-04-HF-mesocosm.csv Harvard Forest field mesocosm experiment hf276-04-HF-mesocosm.csv 9957 182e53e7d4b4f3778170962c87c5fa9c 1 \r\n column , https://harvardforest.fas.harvard.edu/data/p27/hf276/hf276-04-HF-mesocosm.csv date date of the of CO2 efflux measurement YYYY-MM-DD 1 day NA missing value week week of the experiment (e.g. week 1 is the first week of measurements) week of the experiment (e.g. week 1 is the first week of measurements) NA missing value treatment treatment for the mesocosm IM macroinvertebrates, microinvertebrates and microbes present (soil not autoclaved) M microbes and microinvertebrates (soil not autoclaved, no macroinvertebrates added) S salamander, macroinvertebrates, microinvertebrates and microbes present (soil not autoclaved) SI sterile soil with macroinvertebrates only (soil autoclaved) SS sterile soil (soil autoclaved, nothing added) chamber HF ant warming chamber from which the invertebrates originated ant warming chamber invertebrates originated from at HF NA missing value mesocosm unique identifier for each mesocosm unique identifier for each mesocosm NA missing value temp average temperature over the course of the experiment for this mesocosm celsius 0.1 real NA missing value warming mean air temperature delta (difference between chamber and average of 3 ambient reference stations) of Harvard Forest warming chamber over course of experiment celsius 0.000001 real NA missing value sal.pa salamander presence 1 present 0 absent NA missing value invert.pa macroinvertebrates presence 1 present 0 absent NA missing value microbe.pa whether or not soil was autoclaved to remove microbes and microinvertebrates 0 soil autoclaved to remove microbes and microinvertebrates 1 soil not autoclaved NA missing value efflux soil efflux (μmol CO2 mol-1) measured using LI-6400 dimensionless 0.0001 real NA missing value fungal.cover fungus colonization of mesocosm. Qualitative scale from 0 (no visible fungus) to 10 (no bare soil visible) fungus colonization of mesocosm. Qualitative scale from 0 (no visible fungus) to 10 (no bare soil visible) NA missing value invert.mor number of surviving macroinvertebrates/macroinvertebrate number added to mesocosm at start of experiment dimensionless 0.000000001 real NA missing value 179 community short-term measurement https://harvardforest.fas.harvard.edu/exist/apps/datasets/showData.html?id=hf113