DOI: 10.5061/dryad.bg79cnpmr
title: Data from: The wood economics spectrum modulates the positive effects of termite foraging intensity on deadwood invertebrate diversity

description: How populations of ecosystem engineers are both driven by and drive
 biodiversity is poorly known, even less so in detrital subsystems.
 Deadwood plays a key role in maintaining biodiversity and ecosystem
 functions. Wood economics spectrum (WES), which represents the initial
 wood quality through a cluster of wood traits, via afterlife effects might
 affect termite populations and deadwood invertebrate community structure.
 Termites, as ecosystem engineers, exert a significant impact on other
 invertebrate diversity. However, how the WES modulates the effects of
 termite foraging intensity on deadwood invertebrates is unclear. We
 hypothesized that the WES had significant effects on termite foraging
 intensity and deadwood invertebrate abundance and richness. Moreover, the
 WES was hypothesized to modulate the effects of termite foraging intensity
 on deadwood invertebrate diversity. We conducted a wood
 decomposition experiment to test our hypotheses in two subtropical forests
 in China. Logs of 22 tree species with distinct functional traits were
 incubated for 30 months to measure termite foraging intensity (relative
 termite feeding area and the mass of soil materials imported by termites)
 and deadwood invertebrate abundance and richness. We found that from the
 conservative (slow-growing species with low wood quality) to the
 acquisitive (fast-growing species with high wood quality) ends of the wood
 economics spectrum, termite foraging intensity increased. As termite
 foraging activity intensified, the deadwood invertebrate abundance and
 richness increased correspondingly. Moreover, there were significant
 positive relationships between termite foraging intensity and important
 detritivore (Acari, Collembola, and Opisthopora) abundance. In contrast,
 the position along the WES had no direct effect on the abundance and
 richness of deadwood invertebrates besides via termite foraging intensity.
 Synthesis. Our findings showed the pathway through which the WES
 affects deadwood invertebrate diversity. It supported the hypothesis that
 the WES plays a crucial role in shaping the effects of termites as
 ecosystem engineers on the broader invertebrate community. Future studies
 should focus more broadly on whether and how plant traits, via afterlife
 effects on ecosystem engineers, influence invertebrate community
 composition and structure. Such studies will promote our understanding of
 the importance of both plant traits and ecosystem engineer traits for
 ecosystem carbon and nutrient cycling and biodiversity.
description: <strong>Data collected</strong>
 <strong>Invertebrate sampling, identification and
 counting: </strong>In July 2020, 147 logs (22 species) of 35 cm
 length were quickly and carefully put into plastic zip-lock bags in filed
 and taken into the laboratory at Tiantong and Putuo Ecological Station,
 where the samples were stored at 15 °C for subsequent processing. First,
 we put the wood sample in a deep plastic tray and collected larger,
 visibly active invertebrates by hands to prevent escape before processing.
 Then, we sawed the log into two halves, and then we sawed a 2.5 cm thick
 disk at each end of each half log. This way, we collected two disks from
 the middle and end positions. We used one middle disk and one ends disk to
 measure termite foraging intensity, and the other half for mass loss and
 traits analysis (not included in this study). The remaining logs (25-cm
 long) were used to analyze the composition and diversity of the other
 wood-inhabiting invertebrate community.  To collect the
 macroinvertebrates (body width &gt; 2 mm) of logs, firstly we put the
 residual wood sample in a deep plastic tray and separated the bark from
 the wood using chisels or manually. After the bark had been peeled off and
 fragmented into small pieces, we extracted the invertebrates living in the
 bark. The wood remaining after peeling off the bark was split into small
 pieces by an axe, after which we collected the macroinvertebrates by
 hand. The soft xylem with holes from termite consumption was carefully cut
 away with chisels until all soft wood was removed to make sure that all
 invertebrates were collected and transferred them to 30 ml sealed plastic
 bottles with 70% ethanol for identification and counting. If wood under
 the bark was intact and did not show any signs of xylophagy, we would not
 break it for invertebrate searching. The fragmented and soft xylem that we
 cut away were collected to extract the mesoinvertebrates by using a
 modified Tullgren funnel at 35 ℃ for 24 h and collected in 50 ml sealed
 plastic bottles with 70% ethanol solution. All the samples were processed
 within 20 days after collection from the field, i.e. when the
 invertebrates were generally still alive inside the logs.
 Then an electron optical type microscope (DVM6, Leica Germany) at
 a magnification of 35~500× was used for mesoinvertebrate identification
 and counting. All extracted Acari were grouped into Mesostigmata,
 Oribatida and Prostigmata (also including any Astigmata) following Yin
 (1992). All Collembola and adults of other dominant groups were identified
 to family level following Yin (1992, 2000). Juveniles and some rare taxa
 were only identified to the order level due to difficulty of
 identification. Richness was calculated as the number of all other taxa
 (including ants) present per log and abundance was calculated as the total
 number of individuals per log, excluding ants (but conducted a separate
 analysis of termite-ant relationships).
 <strong>Initial wood traits measurement:
 </strong>After we cut all trees into logs, we took the adjacent wood
 disks to the lab for analyses of initial wood traits. We measured the wood
 density (WD), nutrient concentration (N, P concentration), wood dry mass
 content (WDMC), cellulose and lignin content. For measurement of initial
 wood traits (without bark) considered relevant to the WES and to deadwood
 invertebrate preference, each 5 cm disk was stored in a zip-lock plastic
 bag immediately after collection in the field, and kept cool until
 processing. Within 12 h, after the bark had been removed, a subsample was
 cut from each disk to obtain fresh mass and initial volume using
 Archimedes' principle of water displacement. Then wood subsamples
 that had been measured for fresh mass and volume were dried at 75 °C for
 72 h to determine dry mass. Initial wood density
 (g·cm<sup>-3</sup>) was calculated as dry mass per (fresh)
 volume. Initial wood dry matter content was calculated as dry mass/fresh
 mass. Initial dry subsamples were ground in a laboratory mill and passed
 through a 0.15 mm sieve. Thereafter, 0.2 g subsamples were digested using
 concentrated H<sub>2</sub>SO<sub>4 </sub>to
 determine N and P concentrations (mg·g<sup>-1</sup>) on an
 infrared spectrophotometer (Smartchem 200, Alliance, France). Cellulose
 content was determined by anthrone-sulphuric acid colorimetry and
 lignin content was determined by acid detergent fiber (ADF).
 <strong>Measurement of termite feeding area and imported
 soil mass: </strong>After the harvested disks were cleaned
 thoroughly and dried, we first measured the thickness of the disk (for the
 subsequent calculation of disk volume), then digital images of the top and
 bottom surfaces of each disk were taken. The image resolution was enhanced
 (e.g., backgrounds made whiter and brighter, Figure S2a) and areas of the
 remaining wood surface after termite consumption were delineated (e.g.,
 colored white, Figure S2b) using Adobe Photoshop CC19.0. Thereafter Image
 J 1.53c was used to measure the surface area of intact wood. Each wood
 sample had two disks and was measured four times. The average of the
 percentage of (%) termite feeding area and intact wood area of the four
 measurements were used as termite feeding extent due to the relatively
 short length of the wood samples. The damage could still be discerned
 rather clearly in the 30-month samples, but soon after that it became too
 difficult to accurately estimate the damage. We carried
 out a fire experiment to measure the termite imported soil mass as another
 index of termite foraging intensity. The measured soil mass cannot
 represent the activity of all termites, but represent the activity of
 <em>Odontotermes</em> and some
 of <em>Reticulitermes</em> (the dominant species in the study
 area). According to Ulyshen & Wagner (2013), completely burning
 the wood away to leave only the mineral soil behind as ash is a more
 accurate method than washing to isolate soil from decomposed wood and
 collect the soil for quantifying the termite foraging intensity. The fire
 experiment was carried out in the Fire Laboratory located at Hangzhou
 Normal University, Zhejiang province. From September 2020 to January 2021,
 we did a complete combustion of all the wood disks of all 22 species. All
 burns were conducted under a fume hood on a solid fire-resistant plate.
 Before the fire experiment started, the fume hood was turned on and
 ventilated at a constant speed. Prior to each burn, the lab door was
 closed and the fume hood was turned on and the experiment started when the
 room temperature was 21 ± 4 °C. For each burn, we put the wood disk on a
 stainless-steel plate and then put the plate on an electronic heating
 plate. We set the temperature of the electronic heating plate at 300 °C.
 After about 30 min preheating, we ignited the wood by a butane gas torch.
 After ignition, we let the wood burn and smolder naturally until
 completely burning. For wood with low flammability that cannot completely
 burn by itself after ignition, we used a butane gas torch to burn the
 remaining mass completely. <strong>Wood economics
 spectrum (PC1) construction</strong> <strong>:
 </strong>We used a principal component analysis (PCA) to construct
 the wood economics spectrum. Traits in including Carbon, Nitrogen,
 Phosphorus, Lignin, Cellulose, and wood density. Because of the relatively
 high proportion of species variance explained by the first PCA axis, we
 used the PC1 scores to represent a synthetic variable incorporating
 multiple trait variables as an index of
 the<strong> </strong>wood economics spectrum in the subsequent
 analyses. <strong>Dataset  processed:
 </strong>Deadwood invertebrate abundance (the number of individuals)
 data were log-10 transformed and imported soil mass data and termite
 feeding area data were Arcsine-transformed before analysis to better
 achieve normality and linearity. 
description: # The wood economics spectrum modulates the positive effects of termite
 foraging intensity on deadwood invertebrate diversity Ci et al. (2025),
 Data from: **The wood economics spectrum modulates the positive effects of
 termite foraging intensity on deadwood invertebrate diversity** The
 dataset includes the measured data of wood economic spectrums (PC1) based
 on the wood carbon, nitrogen, and phosphorus content, lignin, cellulose
 content, and wood density and relative termite feeding area and termite
 imported soil mass per volume. As well as invertebrate abundance
 (excluding termites and ants) invertebrate richness and the abundance of
 other important detritivores and ants. **Description of the data and file
 structure** * species: species' Latin names - site: incubation site *
 PC1: the values of the first principal component (PC1) after dimension
 reduction of six traits (wood carbon, nitrogen, phosphorus content,
 lignin, cellulose, and wood density) of  wood , representing the wood
 economic spectrum. - Asin_relative_termite_feeding_area:
 Arcsine-transformed relative termite feeding area (%) * Asin_soil
 \_mass_pervolume: Arcsine-transformed imported soil mass per volume by
 termites (g/cm3) - Log_abundance: Abundance of deadwood invertebrates
 (excluding termites and ants) was log10-transformed. * Richness: the
 richness of deadwood invertebrates - Acari_abundance: the abundance of
 Acari * Collembola_abundance: the abundance of Collembola -
 Isopoda_abundance: the abundance of Isopoda * Lumbrieidae_abundance: the
 abundance of Lumbrieidae - Ant_occurrence: the occurrence of ants *
 Ant_abundance: the abundance of ants For additional information, please
 email Hang Ci or Enrong Yan
 ([cihangcaf@163.com](mailto:cihangcaf@163.com),
 [eryan@des.ecnu.edu.cn](mailto:eryan@des.ecnu.edu.cn)).