Deadwood-dependent arthropod community structure and diversity does not change across log type or C. punctulatus abundance

Deadwood-dependent arthropod community structure and diversity does not change across log type or C. punctulatus abundance | Authorea try { document.documentElement.classList.add('js'); } catch (e) { } var _gaq = _gaq || []; _gaq.push(['_setAccount', 'G-8VDV14Y67G']); _gaq.push(['_trackPageview']); (function() { var ga = document.createElement('script'); ga.type = 'text/javascript'; ga.async = true; ga.src = ('https:' == document.location.protocol ? 'https://ssl' : 'http://www') + '.google-analytics.com/ga.js'; var s = document.getElementsByTagName('script')[0]; s.parentNode.insertBefore(ga, s); })(); Skip to main content Preprints Collections Wiley Open Research IET Open Research Ecological Society of Japan All Collections About About Authorea FAQs Contact Us Quick Search anywhere Search for preprint articles, keywords, etc. Search Search ADVANCED SEARCH SCROLL This is a preprint and has not been peer reviewed. Data may be preliminary. 26 March 2026 V1 Latest version Share on Deadwood-dependent arthropod community structure and diversity does not change across log type or C. punctulatus abundance Authors : Taylor Rand and Erin Scott 0000-0003-4581-2523 [email protected] Authors Info & Affiliations https://doi.org/10.22541/au.177451960.06176651/v1 177 views 91 downloads Contents Abstract Information & Authors Metrics & Citations View Options References Figures Tables Media Share Abstract Across a variety of ecosystems, taxonomically distinct primary producers have been shown to support communities that differ in diversity and/or community structure. Additionally, ecosystem engineers and other abundant species have been shown to change the structure and diversity of communities across different ecosystems. Deadwood-dependent (saproxylic) arthropod communities play a crucial role in forest ecosystems, facilitating decomposition of wood and nutrient cycling in forests Decaying deadwood habitats are highly heterogeneous and are able to support large and diverse communities of these deadwood-dependent arthropods. However, our understanding of how these communities differ across different ecosystem factors, such as community composition and characteristics of the wood itself, is limited. We investigated the impacts of two factors on saproxylic arthropod diversity: deadwood type (hardwood vs. softwood) and abundance of Cryptocercus punctulatus, an abundant, social, wood feeding insect. We sampled arthropod communities from 26 logs in the western Appalachian Mountains and identified arthropods to order- and family-level. We observed a large amount of variation in the abundance and diversity of the arthropod taxa sampled, but no effects of log type or C. punctulatus abundance on the arthropod community diversity or community structure. Our study suggests that broad scale deadwood arthropod diversity may not be structured by variation in log type or colonization history of social wood feeding insects, providing clarity about the relationship between two underexplored drivers of diversity within these essential arthropod communities. 1. Introduction Habitat heterogeneity drives community composition by diversifying available resources and microenvironments (Barbour et al 2015). For example, different species of plants have been shown to support different types of arthropod communities (Schaffers et al 2008). Both broad taxonomic differences and subtler genetic differences have been shown to change the diversity and composition of herbivorous arthropods living on plants (Barbour et al 2015, Crutsinger at al 2006, Ferrier et al 2012, Ozanne 1999, Keller et al. 2004). Additionally, abundant or ecologically dominant arthropod species, in particular ecosystem engineers (Jones, Lawton, & Shachack 1994), can change the resources available for other arthropods in the community (Belovsky & Slade 2000, Nicols et al. 2008, Schmidt 2007, Schmitz, Beckerman, & O’Brien 1997), thus further structuring community composition and diversity of other arthropods. The relationship between plant heterogeneity and their associated arthropods has often been studied in the context of living plants. But many forest-dwelling communities rely on deadwood plant resources, with as many as 30% of all forest insect species being deadwood dependent (Speight 1989). Like the habitat provided by living plants, deadwood as a habitat is highly heterogeneous, varying in nutritional content (Meerts 2002), structure (Wiedenhoeft & Miller), size (Henneberg et al. 2021), position (Zumr et al. 2024), decay stage (Zuo et al. 2020), and colonization history (Fukami et al. 2010). However, it is unclear how the specific factors that contribute to the heterogeneity of deadwood influence the composition of the deadwood-dependent arthropod communities. 1.1 Deadwood type Forests containing a mix of softwoods (gymnosperm trees) and hardwoods (angiosperm trees), comprise more than 19 million hectares in Northern U.S. forests and are widespread throughout the rest of the world (Vickers et al. 2021). In these mixedwoods forests, the presence of two distinct wood types creates the opportunity to evaluate the impacts of structural variation on deadwood dependent communities. Hardwoods and softwoods differ in both their anatomical structure and chemical composition. Structurally, softwoods lack the specialized water conduction cells (vessel elements) found in hardwoods [that give them characteristic cross-section pores]. Softwood generally has higher quantities of lignin and other extractives than hardwood (Rowell, Pettersen & Tshabalala, 2012; Kebbi-Benkeder et al., 2014) and lower mineral nutrient (N, K, and Mg) concentrations in their sapwood compared to hardwoods (Meerts 2002), as well as lower rates of decay (Hararuk et al. 2020). These differences could create unique niches and habitats for different deadwood-dwelling arthropods, promoting habitat heterogeneity and altering arthropod community composition. The influence of wood type on community assemblage in deadwood has been investigated across different taxa. Wood type preference has been observed in deadwood-dependent fungal and bacterial assemblages, with several studies documenting differences in species diversity between communities occupying hardwood and softwood logs (Baber et al. 2016; Heilmann-Clausen et al. 2016; Purahong et al. 2018). Observations of differences in arthropod communities across deadwood type, however, show varying trends In some cases, wood type is a strong predictor of Coleopteran and Hymenopteran richness and composition (Seibold et al. 2023; Gossner et al. 2016). The difference in bark traits between tree clades can also substantially influence the structure of early deadwood colonizing arthropod communities (Zuo et al. 2016). In general, however, we lack understanding of how overall arthropod communities encompassing multiple orders differ with wood type. Additionally, the majority of this research into deadwood dependent arthropod communities’ response to wood type differences has been done in European forests, which differ from North American forests in tree species composition, disturbance regimes, and management history (Bouget et al. 2012). It is unclear whether these patterns generalize to eastern North American mixedwoods, especially those of the Appalachian region where deadwood communities remain comparatively understudied (Garrick et al. 2017; Ferro et al. 2012). 1.2 Abundance of Cryptocercus punctulatus Deadwood ecosystems are commonly colonized by large colonies of social wood boring insects, such as termites, beetles, or wood roaches (Hébert 2023) but we do not fully understand how these social insects interact with and influence their cohabitating communities. The presence of abundant social insects within deadwood could result in positive diversity outcomes for the broader deadwood dependent arthropod community by increasing heterogeneity and microhabitat availability through excavation of cavities and tunnels. Some existing research supports this idea. For example, microhabitats created by wood boring beetles have been found to support and increase the diversity in arboreal ants (Priest et al. 2021), trophic isopods (Tuo et al. 2024), multi-taxa arthropod assemblages (Buse et al. 2008), and even reptiles (Gottfried et al. 2019) living in deadwood. The development of microhabitats by termites, too, has been found to increase cohabitating invertebrate community diversity (Ci et al. 2025). However, colony defense or competition between social insects and their cohabitants could also result in negative impacts on arthropod community diversity in deadwood. This idea is supported by research in post-burn habitats, showing that in a post-burn context, termite abundance can have an either neutral or negative relationship with cohabitating beetle community diversity (Ulyshen et al. 2020). Cryptocercus punctulatus is a common deadwood-dwelling social insect in Appalachian forests (Fig. 1). C. punctulatus is a sub-social wood roach species that builds complex galleries within fallen logs and occupies a diversity of log types (Nalepa 2020). C. punctulatus adults are physically large for an insect (typically over 2.5 cm long and weighing up to 1g). Mate pairs can have broods of up to 75 nymphs, which take over a year to mature (Nalepa, 1990). C. punctulatus galleries alter the physical structure and available habitat space of the logs where they occur, and the large volume of excrement present in the galleries could alter chemical and nutritional properties or microbial colonization of the log. Currently, we do not understand of how presence and abundance of C. punctulatus affects deadwood arthropod communities, and whether patterns of association match the ones observed in other social deadwood dwelling insects. 1.3 Our Questions Many studies on the diversity and community structure of deadwood dependent arthropods focus on specific taxonomic groups like termites, beetles, or spiders (Muller et al. 2020; Wende et al. 2017; Økland et al. 1996; Zumr et al. 2024; Ci et al. 2025). While some recent studies have expanded diversity analyses to include multiple taxa (Garrick et. al 2019, Andringa et al. 2019; Vogelfanger et al. 2025) there is a remaining need for studies that link variation in deadwood habitats to variation in overall arthropod community structure and diversity. In this study, we focus on two factors that may create heterogeneity in deadwood habitats: A) deadwood type, B) the abundance of an abundant social, wood feeding insect, C. punctulatus . We investigate how arthropod diversity and community structure differs across each of these factors. 2. Methods 2.1 Sampling Sites 26 logs were sampled from 3 montane mixedwoods sites in Giles County, Virginia: White Rock Branch (37.2549, 80.3005, elev. 890m), Jungle Trail (37.3677, -80.5334, elev. 1200m), and Pond Drain (37.2144, 80.3220, elev. 1256m). All sites are positioned near Mountain Lake Biological Station (MLBS), with Jungle Trail ~1.4 km to the northwest of MLBS, White Rock Branch ~ 6.5 km to the northeast, and Pond Drain ~2km to the southwest. All three sites experience cool climate conditions (summer daytime temperatures averaging around 21°C, nighttime temperatures ranging from 4-15°C). The tree composition at all three sites included hardwoods such as various species of oak ( Quercus , especially Quercus alba ), maple ( Acer , especially Acer rubrum ), cherry ( Prunus sp. ), and tulip poplar ( Liriodendron ), with intermittent stands of Eastern hemlock ( Tsuga canadensis ). 2.2 Sampling and Identification 8-10 logs were sampled per site. Sampling began at the thick end of each log, and continued down the log until no new arthropod taxa had been found for 15 minutes of sampling time. All logs sampled were at intermediate to late decay stage, designated in accordance with the USDA Forest Service’s decay stage classifications (Russell et al. 2013). To account for natural disparities in deadwood size between tree species and tree types, we measured total length of the log and length sampled, as well as diameter at the base and tip of the log. We used these measurements to calculate volume sampled for each log. Logs were taken apart using a hammer and chisel. Arthropods were sampled from three sections of the log: beneath the bark, within the sapwood, and within C. punctulatus tunnels that go deep into the heartwood. When C. punctulatus galleries were not present, the heartwood was too difficult to access and thus was not sampled. Trash bags were tucked around and under logs to prevent loss of samples in the surrounding leaf litter. Wood chips were piled on a separate trash bag during sampling. After the log had been sampled, we surveyed discarded wood chips for approximately 15 minutes and collected any missed arthropods. Only arthropods >2 mm in length were sampled. For arthropod taxa which could be easily identified on site, abundance was recorded during sampling. Voucher samples for these taxa were collected and preserved for future reference. Any specimens which could not be identified on site were collected, preserved, photographed, and identified in the lab. For all taxa except ants, abundance was recorded as the exact number of individual arthropods observed for that taxonomic group. In the specific case of ants (family Formicidae), abundance was estimated at 100 individuals per nest, as ants were generally too abundant to count when present. The taxonomic level to which we were able to identify samples varied between arthropod clades. Taxa from classes Arachnida, Chilopoda, and Diplopoda were identified only to the taxonomic rank of order, due to a lack of identification resources for these groups. All insect taxa were identified to the rank of family or superfamily using dichotomous keys (Triplehorn et al. 2005, Chu 2012) or assigned a unique morphotaxon name if exact family name was unknown. 2.3 Data Analysis Calculating log volume After sampling, we calculated total volume and volume sampled for each log using length, tip diameter, and base diameter measurements that we had taken on site. Total volume was calculated by modelling logs as conical frustums (i.e. a cone without apex) using\(V=\frac{\text{πh}}{3}\left(r^{2}+\text{rR}+R^{2}\right)\) where\(r\) is the radius at the log tip, \(R\) is the radius at the log base, and \(h\) is the log length. A conical frustum shape was chosen to represent the complex log shape instead of a cylinder or simple cone, as it better reflected the logs’ tapered shape. Volume sampled was approximated using a simple cylindrical model (\(V=2\pi R^{2}h\)) based on the base radius and length of the log sampled, providing a realistic estimate of the volume represented by the sections sampled. Using a linear regression model, we determined there was no relationship between volume sampled, length sampled, total log length nor total log volume and arthropod diversity or community structure across the data set; we did not then include it as a parameter in our models. Describing log type and C. punctulatus abundance Log type was first determined in the field using the presence of branch nubs on the logs (softwood logs retain sturdy branch stubs even after the log has significantly decayed, while hardwood logs do not). Wood samples from each log were additionally collected, and log type was confirmed in the lab by examining the wood grain under a microscope (identifying the presence of hardwood characteristic rays and softwood characteristic pores). The absence of bark and other identifiable features on intermediate to late decay logs made species-level identification of logs unfeasible. The number of softwood and hardwood logs was balanced across sites, with similar numbers of hard and softwood sampled at each site, and similar numbers of total logs sampled across sites (Table A3). C. punctulatus abundance for each log was delineated as zero (no adults found), low (1-3 adults found), medium (4-10 adults found), or high (11 or more adults found) These grouping choices were made a priori based on the physical effects we perceived each category typically had on the logs they occupied during practice sampling (i.e. amount of feces present, proportion of log with galleries, length and density of tunnels). We did not consider the abundance of C. punctulatus nymphs, though most logs with at least 1-3 adults had nymphs present as well. As we wanted to assess the effects of C. punctulatus abundance and presence on the rest of the arthropod community, we did not include C. punctulatus counts in our calculations of arthropod community diversity or community structure. Due to the a priori grouping choices and our inability to determine how abundant C. punctulatus were within a log before sampling, the number of logs in each category of C. punctulatus abundance had some imbalance across sites. We sampled no logs with medium C. punctulatus abundance sampled at Jungle Trail and no logs with high C. punctulatus abundance sampled at White Rock Branch (Table A4). Arthropod diversity (Simpson’s diversity and species richness) Data analysis was done in RStudio using R version 4.4.2. For each log, richness was calculated as the number of arthropod taxa present. Simpson’s diversity index was calculated using the vegan package (version 2.7-0; Oksanen J, et al. 2025). The effects of wood type and C. punctulatus abundance on richness and Simpson’s diversity (Simpson’s) were evaluated using generalized linear mixed models via the car package (version 3.1-3; Fox J, Weisberg S, 2019). Separate models were used for analyses of Simpson’s and richness. Each model included log type, C. punctulatus abundance and site as fixed effects. Effects were modeled as diversity ~ C. punctulatus abundance + log type + site The contribution of main effects was tested using a type II ANOVA, which evaluates each predictor after accounting for all other main effects in the GLM. Model assumptions were tested using the DHARMa package (version 0.4.7; Hartig F, 2024). P-values for level comparisons were generated using the emmeans package (version 1.11.2-8, R Lenth, 2025) and estimated marginal means were plotted for visualization. Using a separate linear regression model, we determined there was no relationship between volume sampled, length sampled, total log length or total log volume and arthropod diversity across the data set, therefore we did not include any of these variables as effects in our diversity GLMs. Community structure We chose Bray-Curtis community dissimilarity to represent differences in the ecological structure of arthropod communities in our study, as this measure of beta diversity describes community composition with sensitivity to both abundance and presence absence of taxa. Bray-Curtis distances representing dissimilarity in arthropod communities were calculated for each pair of logs using the vegan package (version 2.6.10; Oksanen J, et al. 2025) A 2D ordination NMDS ordination was generated to visually represent Bray-Curtis distances between arthropod communities in each respective log. The effects of wood type and C. punctulatus abundance on arthropod community structure, as measured by Bray-Curtis, were evaluated using a PERMANOVA. The PERMANOVA tested whether dissimilarity between logs within treatments was greater than the dissimilarity between logs among treatments, thus evaluating whether community structure differed across treatment in our study. The PERMANOVA model included log type, C. punctulatus abundance, and site as effects. Effects were modeled as Dissimilarity ~ C. punctulatus abundance + log type + site. As with the alpha diversity analysis, we determined using separate PERMANOVAs that there was no relationship between volume sampled, length sampled, total log length or total log volume and Bray-Curtis distances across the data set, therefore we did not include any of these variables as effects in our main PERMANOVA. The contribution of main effects was tested using a marginal analysis of variance, which evaluates each predictor after accounting for all other main effects in the PERMANOVA. We used the betadisper and permutest functions to evaluate whether average distance to the ordination centroid differed across levels of log type, site, and C. punctulatus abundance. For effects with non-homogenous distances to the centroid, we used a Tukey test to determine which specific groups differed from each other. 3. Results 3.1 Arthropod community variation within our study Over a 1-month sampling period, 1,596 arthropods were sampled and identified from 33 taxonomic groups, composed of 11 orders from classes Arachnida, Diplopoda, Chilopoda, and Malacostraca and 28 families (or superfamilies) from class Insecta. The most abundant taxon sampled was family Formicidae (ants). Orders Coleoptera and Aranea (beetles and spiders) were the second and third most abundant arthropod groups, respectively. Arthropod richness varied widely, with logs possessing anywhere from 8 to 40 unique taxa. Taxa highly represented in the dataset include beetles (predominately Superfamily Scarabaeoidea, followed by Families Elateridae, Carabidae, Lucanidae, and Staphylinidae), centipedes (Orders Scolopendromorpha, Geophilomorpha, Lithobiomorpha), and millipedes (Orders Julidae, Chordeumatida, and Polydesmida). We did not observe any Bess beetles (Family Passalidae) or termites (Infraorder Isoptera), two social insect groups commonly associated with deadwood. Rather, either ants (family Formicidae) or C. punctulatus were often the most abundant arthropods in each log 3.2 No effect of log type or C. punctulatus abundance on arthropod diversity We used two separate GLMs to evaluate the effects of log type and C. punctulatus abundance on arthropod richness and Simpson’s diversity respectively. We found no effect of log type on Simpson’s diversity (Figure 2a, Table A1; p = 0.397) or richness (Figure 2b, Table A2, p = 0.769). This result suggests that neither the number of arthropod taxa present nor the abundance distribution of those taxa is predicted by log type in the habitats we sampled. Similarly, neither Simpson’s diversity (Figure 3a, Table A1; p = 0.315) nor richness (Figure 3b, Table A2; p = 0.432) differed across levels of C. punctulatus abundance. This result suggests that neither the counts nor relative abundance distributions of arthropod taxa is influenced by C. punctulatus abundance in our habitats. 3.3 No effect of log type or C. punctulatus abundance on arthropod community structure We used a PERMANOVA to evaluate whether arthropod community structure, quantified using Bray–Curtis dissimilarity among log communities, differed across log types or levels of C. punctulatus abundance. Arthropod community structure did not differ by log type (Figure 4a, Table A5; p = 0.43) or by C. punctulatus abundance (Figure 4b, Table A5; p = 0.88) across the logs sampled in our experiment. Because Bray–Curtis dissimilarity reflects differences in both taxon presence–absence and relative abundance, these results indicate that neither log type nor C. punctulatus abundance was associated with differences in arthropod abundance or community composition. We also used dispersion testing to evaluate whether arthropod communities differed in how variable they were within groups defined by log type or C. punctulatus abundance (in contrast to the PERMANOVA, which evaluated whether average community composition differed between groups). Dispersion was quantified as the average distance of each log’s arthropod community to the centroid of its group in Bray–Curtis ordination space, such that smaller distances indicate greater similarity among logs within the same group. Dispersion did not differ between hardwood and softwood logs (Table A6, p = 0.83), indicating comparable levels of within-type community variability. In contrast, dispersion differed across levels of C. punctulatus abundance. Specifically, logs with high C. punctulatus abundance were less dispersed than logs with zero C. punctulatus abundance (Figure 5, Table A6; p = 0.043), indicating that arthropod communities were more similar to one another within the high-abundance category than within the zero-abundance category, even though average community composition did not differ between abundance levels. 4. Discussion We did not find significant effects of either log type or C. punctulatus abundance on arthropod diversity (richness and Simpson’s diversity). We also found that neither log type nor C. punctulatus abundance predicted arthropod community structure. These results suggest that log type and C. punctulatus abundance are not primary drivers of arthropod diversity, richness, relative abundance, or composition in the habitats we sampled. However, we also found more similarity between logs within the high-abundance category compared to logs in the low abundance category. This indicates that while C. punctulatus abundance does not change average community structure, it can influence variability in community structure. Our prediction that that arthropod communities would differ across different log types was based on an expectation that structural and chemical differences between hard- and softwoods could represent different ecological niches and thus create ecologically distinct habitats for arthropods. Our prediction was additionally based on evidence that log type is an important predictor of community ecology for other deadwood colonists, such as bacteria and fungi, leading us to expect a similar pattern in arthropods. Our results, however, did not support this prediction. It is possible that broad-scale arthropod community differences across site could have masked any more subtle differences in arthropod community ecology across log type in our study. This idea is supported by our data. Site was a significant predictor of both Simpson’s diversity and community structure in our analysis, explaining a large percentage (~20%, table citation) of variation in community structure. Though we accounted for site in our calculations log type contributions, these site-level differences may still have reduced power to detect variation in Simpson’s diversity and community composition in our analyses. Richness, on the other hand, was not influenced by site, and thus lack of differences in richness across log type cannot be attributed to site-level effects. Another possible explanation for our results is that other deadwood features are more important for arthropod colonization than wood type (hardwood vs. softwood). Previous work assessing multi-taxa diversity in deadwood suggests that host log type may be a less important contributor to arthropod diversity than other environmental factors, particularly decay stage. Studies conducted on specific insect groups in temperate European and North American forests support this pattern. Irmler et al. (1996) found that wood age class had a stronger influence than tree species on the diversity and abundance of saproxylic beetles and midges in northern Germany. Similarly, Müller et al. (2020) identified decay stage alongside microclimate as the primary drivers of beetle and spider diversity in German forests. Wende et al. (2017) further demonstrated that tree-species specialization within xylophagous beetle interaction networks declined with increasing successional age of deadwood, indicating reduced host specificity over time. In contrast, Varady-Szabo and Buddle (2006) reported generally weak effects of both wood type and decay stage on spider assemblages in Quebec maple forests, although spider diversity was higher in less decayed logs than in advanced stages. More recent multi-taxa studies reinforce the primacy of decay stage over tree identity. Using metabarcoding across more than 4,000 invertebrate taxa in long term deadwood experimental site in German forests, Vogelfanger et al. (2025) found that time since tree death was the strongest determinant of invertebrate community composition and richness whereas tree species identity had comparatively minor effects. Likewise, Andringa et al. (2019) showed that although different tree species and decay stages can support distinct assemblages, variation among decay stages explained substantially more community turnover than tree species alone in the Netherland forest study site. Taxon-specific preferences for wood type appear most apparent in earlier stages of decay, a trend often observed in communities of deadwood-dwelling beetles (Jonsell et al. 2007; Stokland et al. 2012). Wende et al. (2017) demonstrated that specialization in deadwood-dwelling beetles is strongest during the early successional stages of deadwood decomposition and weaken as deadwood ages. As decomposition progresses, structural properties, fungal colonization, and nutrient profiles increasingly converge across tree species (Zuo et al. 2016). Patterns of community homogenization and a decrease in tree-species or tree character specialization is seen in deadwood-dependent Diptera (Mlynarek et al. 2018) as well as in Isopoda, Chilopoda, Myriapoda (Zuo et al. 2020). Because we sampled only intermediate- to late-stage logs, our findings may therefore reflect this homogenization process, in which decay dynamics override initial differences associated with hardwood versus softwood substrates. It is also possible that the absence of differences in arthropod community ecology across log type could be attributed to the abundance of higher tropic-level taxa that use deadwood primarily as habitat as opposed to a food resource. Tertiary consumers like spiders (Araneae) and centipedes (Chilopoda) were high in abundance within our data set and are commonly associated with deadwood ecosystems (Garrick et al. 2019). Because taxa from higher trophic levels depend on deadwood as habitat from which to feed on deadwood-dwelling fungi or fauna, tree species could be less important in structuring interaction networks at higher tropic levels (Wende et al. 2017). Similarly, ants (Formicidae), which comprised a large portion of total arthropod abundance in our samples, function in many habitats as opportunistic foragers with broad diets. If abundant taxa within our communities respond primarily to prey availability or surrounding resource availability rather than specific deadwood characteristics, this could explain limited association between log type and arthropod community structure in our study. Our prediction that C. punctulatus abundance would influence the ecology of co-occurring arthropod communities was based on the expectation that their physical manipulation of the deadwood environment would create distinct environments within log habitats or increase diversity of microhabitats available within the log. Our results showing that variability in community structure is greater in logs with no C. punctulatus presence than in logs with high C. punctulatus abundance provides some support for this idea. It is possible that a high density of C. punctulatus reduces niche availability with a log habitat, thus decreasing the potential for different kinds of communities to live there and homogenizing community structure of co-occurring arthropods within logs highly occupied by C. punctulatus . Though there is some imbalance in the distribution of levels of C. punctulatus abundance across sites in our sampling design, it is unlikely that variability across sites is being confounded with variability across levels of C. punctulatus abundance, as variability in community structure did not differ across sites. Despite our findings of differing variability in community structure across levels of C. punctulatus abundance, we still see no difference in average community structure across levels of C. punctulatus abundance. We also see no difference in richness or Simpson’s diversity across levels of C. punctulatus abundance, in contrast to our original hypothesis. As with our log type results, the strong effect of site may mask more subtle effects of C. punctulatus abundance on Simpson’s diversity and community structure (though not richness), providing one potential explanation for the lack of effects we observed. Other abundant arthropods within deadwood ecosystems modify their deadwood substrates in ways that result in ecological changes to co-occurring arthropod communities. Many wood-boring beetles are xylophagous during development, excavating extensive galleries that facilitate colonization by secondary wood-borers and create microhabitats associated with sap, frass, and decomposing tissues (Graham 1925; Feller and Mathis 1997; Larson and Harman 2003). Termites likewise act as ecosystem engineers, as their feeding activity and associated soil imports can strongly influence nutrient dynamics and have been shown to explain patterns of arthropod biodiversity within deadwood (Ulyshen 2014; Ci et al.). Other taxa, such as ambrosia beetles, carpenter ants ( Camponotus spp.), and wood-nesting bees, excavate galleries for fungal cultivation or nesting, further altering internal log structure (Ulyshen 2014; Rossi and Feldhaar 2019). In our study, wood-boring beetles (including Cerambycidae and Elateridae) and carpenter ants were present. These co-occurring taxa may independently generate substantial structural and microenvironmental change. Effects of other ecosystem engineers could buffer or obscure the detectable effects of C. punctulatus abundance on arthropod community structure. This idea is supported by our results, which show that a large percentage of variation in community structure in our study (~80%) is explained by factors other than log type, C. punctulatus abundance, or site. One of these factors could be structural change introduced by other taxa. Studies describing relationships between other social, abundant, wood-dwelling insects and general arthropod ecology within deadwood ecosystems provide important points of comparison with ours. Termites, the closest relatives of Cryptocercus , are abundant, social insects that substantially alter the structural composition and microhabitat conditions of their woody substrates through feeding and tunneling (Ulyshen 2014). Tunneling and foraging activity in termites have been positively associated with increased cohabiting arthropod richness and abundance in some systems (Ci et al. 2025). However, the influence of termites on co-occurring arthropods can be highly variable and taxon specific. Ulyshen et. al (2020) found in a post-burn habitat, the presence of termites had a largely neutral association effects on co-occurring beetle and ant species: out of 197 total co-occurrence pairings, 96.09% were neutral versus 3.55% negative and 0.56% positive. These results indicate that even when termites substantially modify log structure, their effects on broader insect communities may be limited to particularly sensitive taxa rather than producing broad scale shifts in community composition. Similarly, although C. punctulatus is capable of structural modification of deadwood, we did not detect a relationship between its abundance and overall arthropod community structure. The ecological influence of C. punctulatus may therefore be context dependent, varying across different habitats and regions, or may be restricted to particular taxa that we did not analyze in isolation. More broadly, ecological research outside of deadwood systems indicates that the effects of dominant or ecosystem-engineering taxa on community diversity in general are highly variable and context dependent. While ecosystem engineers are globally associated with increases in local species richness, the magnitude and direction of these effects vary across ecosystems, latitudes, and engineer types, with weaker or inconsistent richness effects reported in higher-latitude systems and temperate forests similar to the mountainous temperate forests that makes up our study sites. (Romero et al. 2014). This difference might be attributed to the fact that higher-elevation temperate forests often support smaller regional species diversity and weaker density-dependent interactions than those of the tropics and tropical forests, potentially limiting the extent to which habitat engineering can translate into species diversity increases (Romero et al. 2014). As our study was done in a high latitude, temperate forest environment, our results showing the relationship between C. punctulatus and deadwood-dwelling arthropod diversity could be consistent with this trend. It is important to acknowledge how our sampling design may constrain our results. The dominant softwood species in our study was Eastern hemlock, while a more diverse set of hardwood species, including oak ( Quercus ), maple ( Acer ), and cherry ( Prunus ), were sampled. Hemlock may provide distinct ecological niches that could influence community composition differently than other softwood species, and thus not be representative of mixedwoods forests with more softwood diversity represented. A similar study in an area with more softwood diversity could yield different results. It is also possible arthropod communities are more responsive to deadwood species (i.e., oak vs. maple) versus wood type (hardwood vs. softwood) as defined in this study. Additionally, our dataset contained relatively few logs with medium or high C. punctulatus abundance, which may have limited our ability to detect significant relationships. We also sampled exclusively during mid-summer. Many wood-dwelling arthropod species only use deadwood at specific life stages which are likely seasonal, so more exhaustive sampling across seasons could give a more comprehensive snapshot of saproxylic arthropod communities. 5. Conclusion Our investigation into multi-taxa ecological trends in deadwood shows that neither Cryptocercus punctulatus abundance nor log type significantly structures deadwood-dependent arthropod community ecology in late-stage decay logs within mixedwoods forests of the southeastern Appalachian mountains. These results reveal that deadwood-dependent arthropod communities can maintain similar broad taxonomic diversity, richness, and community structure across environmental and ecological variations in deadwood. Deadwood offers an important habitat and nutritional resource for a large and diverse amount of arthropod forest taxa (Ulyshen & Sobotnik 2018, Garrick et al. 2019), whose populations are threatened by fragmentation of habitat and removal of deadwood in managed forests (IUCN 2025, Tsikas & Karanikola 2022, Siitonen 2001). Because of the essential roles deadwood-dependent arthropods play in wood decomposition, nutrient cycling, and habitat creation for other forest organisms (Ulyshen 2018, Hardersen & Zapponi 2018), it is important to investigate potential drivers of diversity and community composition to better understand broader ecological dynamics and support conservation efforts. Our study provides clarity about the relationship between two underexplored drivers of diversity within these essential arthropod communities. Acknowledgements We thank Margaret Cumberland at the National Ecological Observatory Network (NEON) for assistance with deadwood identification. We thank Dr. Jessamyn Manson (University of Virginia, School of Arts and Sciences) for advice on sampling design and for reviewing an early draft of this manuscript. We also thank Joseph Mullica for providing photographs of Cryptocercus punctulatus . Finally, we thank the National Science Foundation Research Experience for Undergraduates (NSF REU) program for funding this work. Data Availability Statement Our code and associated README files are available on GitHub under repository name Rand_Scott_deadwood (https://github.com/taylorrand/Rand_Scott_deadwood). Within the repository is an R file (finaldata_3_7_26.R), containing the code used to analyze our data and draw our conclusions, and an RData file (finaldata_3_7_26.RData) which contains all objects and variables generated by the R file code. A README file is also available as a guide for the R code and a description of the data and analysis methods. References 1. Andringa, J. I., Zuo, J., Berg, M. 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Ecosystems , 24 (4), 926–938. https://doi.org/10.1007/s10021-020-00558-9 FIGURES, TABLES, AND APPENDIX Figure 1. Figure 1. Adult Cryptocercus punctulatus , Figure 2. Figure 2. Estimated marginal means of Simpson’s diversity index (Simpson’s) for A) deadwood arthropod communities from hardwood and softwood logs and B) communities cohabitating with various abundances of Cryptocercus punctulatus , grouped into categories according to individual adult roaches found in the logs: zero (0), low (1-2), medium (4-10), and high (11+). Points represent model-estimated marginal means derived from generalized linear models (diversity index ~ (log type + C. punctulatus abundance) + site, family = gaussian), with error bars indicating 95% confidence intervals. Numbers above points represent sample size for each wood type. Figure 3. Figure 3. Estimated marginal means of taxonomic richness for A) deadwood arthropod communities from hardwood and softwood logs and B) communities cohabitating with various abundances of Cryptocercus punctulatus , grouped into categories according to individual adult roaches found in the logs: zero (0), low (1-2), medium (4-10), and high (11+). Points represent model-estimated marginal means derived from generalized linear models (diversity index ~ (log type + C. punctulatus abundance) + site, family = gaussian), with error bars indicating 95% confidence intervals. Numbers above points represent sample size for each wood type. Chi^2 df Pr(>Chisq) type 0.556 1 0.456 C. punctulatus abundance 3.55 3 0.315 site 7.02 2 0.030* Chi^2 df Pr(>Chisq) type 0.001 1 0.970 C. punctulatus abundance 2.75 3 0.432 site 2.55 2 0.280 Site Hardwood Softwood Jungle Trail (JT) 3 5 Pond Drain (PD) 5 5 White Rock Branch (WR) 4 4 Site Zero (0) Low (1-3) Medium (4-10) High (11+) Jungle Trail (JT) 2 4 0 2 Pond Drain (PD) 4 1 2 3 White Rock Branch (WR) 3 3 2 0 Effect Sum of Sqs R2 F p C. punctulatus abundance 0.41 0.067 0.63 0.88 Log type 0.2 0.032 0.91 0.43 site 1.2 0.19 2.7 0.014* Residual 4.1 0.67 Contrast Difference Lower CI Upper CI p Low-Zero -0.00551 -0.104 0.0933 0.999 Medium-Zero -0.0359 -0.158 0.0863 0.846 High-Zero -0.117 -0.23 -0.00317 0.0425* Medium-Low -0.0304 -0.155 0.0941 0.904 High-Low -0.111 -0.227 0.00484 0.0636 High-Medium -0.0807 -0.217 0.0557 0.377 Information & Authors Information Version history V1 Version 1 26 March 2026 Copyright This work is licensed under a Non Exclusive No Reuse License. Keywords community ecology ecological experiment ecosystem invertebrate terrestrial Authors Affiliations Taylor Rand University of South Florida View all articles by this author Erin Scott 0000-0003-4581-2523 [email protected] University of Virginia View all articles by this author Metrics & Citations Metrics Article Usage 177 views 91 downloads .FvxKWukQNSOunydq8rnd { width: 100px; } Citations Download citation Taylor Rand, Erin Scott. Deadwood-dependent arthropod community structure and diversity does not change across log type or C. punctulatus abundance. Authorea . 26 March 2026. DOI: https://doi.org/10.22541/au.177451960.06176651/v1 If you have the appropriate software installed, you can download article citation data to the citation manager of your choice. Simply select your manager software from the list below and click Download. For more information or tips please see 'Downloading to a citation manager' in the Help menu . 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