Forests 2012, 3(3), 632-652; doi:10.3390/f3030632

Effects of Small-Scale Dead Wood Additions on Beetles in Southeastern U.S. Pine Forests
Kier D. Klepzig 1,*, Michael L. Ferro 2, Michael D. Ulyshen 3, Matthew L. Gimmel 4, Jolie B. Mahfouz 5, Allan E. Tiarks 5 and Chris E. Carlton 2
USDA Forest Service, Southern Research Station, 200 WT Weaver Blvd, Asheville, NC 28804, USA
Department of Entomology, Louisiana State University, 404 Life Sciences Building, Baton Rouge, LA 70803-1710, USA; Email: (M.L.F.); (C.E.C.)
USDA Forest Service, Southern Research Station, 201 Lincoln Green, Starkville, MS 39759, USA; Email:
Department of Ecology & Evolutionary Biology, University of Kansas, Lawrence, KS 66045, USA; Email:
USDA Forest Service, Southern Research Station, 2500 Shreveport Hwy, Pineville, LA 71360, USA; Email: (J.B.M.); (A.E.T.)
Author to whom correspondence should be addressed; Email:; Tel.: +1-828-257-4307; Fax: +1-828-257-4313.
Received: 19 June 2012; in revised form: 14 July 2012 / Accepted: 25 July 2012 /
Published: 15 August 2012


: Pitfall traps were used to sample beetles (Coleoptera) in plots with or without inputs of dead loblolly pine (Pinus taeda L.) wood at four locations (Louisiana, Mississippi, North Carolina and Texas) on the coastal plain of the southeastern United States. The plots were established in 1998 and sampling took place in 1998, 1999, and 2002 (only 1998 for North Carolina). Overall, beetles were more species rich, abundant and diverse in dead wood addition plots than in reference plots. While these differences were greatest in 1998 and lessened thereafter, they were not found to be significant in 1998 due largely to interactions between location and treatment. Specifically, the results from North Carolina were inconsistent with those from the other three locations. When these data were excluded from the analyses, the differences in overall beetle richness for 1998 became statistically significant. Beetle diversity was significantly higher in the dead wood plots in 1999 but by 2002 there were no differences between dead wood added and control plots. The positive influence of dead wood additions on the beetle community can be largely attributed to the saproxylic fauna (species dependent on dead wood, such), which, when analyzed separately, were significantly more species rich and diverse in dead wood plots in 1998 and 1999. Ground beetles (Carabidae) and other species, by contrast, were not significantly affected. These results suggest manipulations of dead wood in pine forests have variable effects on beetles according to life history characteristics.
arthropods; biodiversity; coarse woody debris; epigeic; insects; slash

1. Introduction

Logs and other woody debris on the forest floor are an essential habitat for a wide range of saproxylic arthropods [1,2,3], which are directly or indirectly dependent on dead and dying wood [4]. These include not only phloem and wood feeders but also their predators and species associated with wood-decaying fungi. Many non-saproxylic ground-dwelling arthropods benefit from dead wood as well. For instance, a number of studies have shown litter-dwelling arthropods and other invertebrates to be more numerous immediately next to dead wood than short distances away from it [5,6,7,8,9,10,11,12,13,14,15,16,17]. These results can be attributed variously to dead wood providing a relatively stable source of moisture [18,19,20], shelter from predators or adverse weather conditions, and an abundance of prey or other food items.

With demand for forest products continuing to rise (especially now with a growing interest in biofuel production, see [21,22]), understanding the role of dead wood in maintaining biodiversity, tree productivity and long-term forest sustainability is becoming more important in forest management. Of particular value are studies investigating the effects of dead wood manipulations on these interests. Several manipulative studies have demonstrated the positive relationship between dead wood and ground-dwelling arthropod diversity [23,24,25], but the question is far from resolved and has received little attention in many forest types.

The forests of the southeastern United States are among the most productive in North America with loblolly pine (Pinus taeda L.) being the single most important timber species in the region. One of the few efforts to measure arthropod response to dead wood manipulations made in loblolly pine plantations began on the Savannah River Site in South Carolina in 1996. In the first phase of that study, plots in which all dead wood was removed annually were compared to reference plots from which no dead wood was removed. After pitfall-sampling ground-dwelling arthropods in those plots for five years, Hanula et al. [26] found no differences in overall abundance or morphospecies richness. Overall arthropod diversity and evenness were significantly lower in removal plots than in reference plots, however, and several families differed in abundance between the two treatments. These differences were observed only in the first two full years of sampling, however. In the second phase of the study, beginning in 2001, arthropods were sampled in these same plots for four more years. Two new treatments were added in which sampling also took place; one involving major inputs of logs to the forest floor and the other involving major inputs of dead standing trees. Contrary to expectations, but largely consistent with the first phase of the study, there were no differences in total ground-dwelling arthropod abundance, richness, diversity or composition among treatments [27]. Only ground beetles (family Carabidae) exhibited a significant association with dead wood.

During 1998, a study was installed on four widely-separated USDA Long Term Soil Productivity (LTSP) sites in the Southeastern United States to better understand the importance of dead wood to nutrient cycling and biodiversity in pine forests. The responses of beetles, the most diverse arthropods on the forest floor, are reported here.

2. Materials and Methods

This study took place at the following four LTSP sites: Croatan National Forest, Craven Country, NC, USA; DeSoto National Forest, Jones Country, MS, USA; Kisatchie National Forest, Rapides Parish, LA, USA; Davy Crockett National Forest, Trinity Country, TX, USA. The forests were dominated by 50–60 yr pine [loblolly in LA and NC, slash (P. elliottii Engelm.) in MS, and loblolly and shortleaf (P. echinata Mill.) in TX] with an understory consisting of oak (Quercus sp.), hickory (Carya sp.), and sweetgum (Liquidambar styraciflua L.). For this experiment, the coarse woody debris was added to remnant mature pine stand, adjacent to the LTSP treatment plots. At each site, three of the mature stand plots were used as controls and contained only the existing woody debris already present or that which fell during the study. A 30 × 45 m rectangular area at each site was divided into six 15 × 15 m plots. The dead wood added to all of the plots came from the same 43-year-old loblolly pine stand in the Palustris Experimental Forest, Louisiana. Sections of wood were removed from the main trunk (i.e., “logs”, 30–35 cm dbh, 1 m long), limbs (2.5–10 cm mid-length diam, 0.5 m long), and twigs (0.5–1.5 cm diam, 0.25 m long) of loblolly pine trees felled for this purpose. Ten logs, 8 limbs, and 24 twigs were distributed across the three randomly selected dead wood plots at each site in April and May 1998. At each given site, the material was added all at once, on one date during the time frame mentioned above. At all sites, the amount of existing coarse woody debris and the vegetation on the ground were roughly the same. Our additions of material impacted the sites in a very similar manner, adding the above material to the baseline of that already onsite.

One pitfall trap (modified from [28]) was installed in each plot to sample ground-dwelling beetles. At each site, the trap was placed arbitrarily within the plot, in the immediate area of the added dead wood. Each trap consisted of a 15-cm-diameter plastic funnel fastened beneath a hole in the center of a 30 × 30 cm section of ~1.9 cm-thick plywood with bevelled edges. This assembly was positioned over a 2-liter plastic container, half-filled with a 1:1 mixture of propylene glycol and 95% ethanol, buried at ground level. Samples were collected approximately biweekly from May through September at all locations in 1998 and at all locations except for North Carolina in 1999. Additional samples were collected in all locations except for North Carolina in April and May of 2002.

Specimens not identified below family level (117 individuals were only identified to family level) were excluded from the dataset. After pooling all sampling periods for a given year, analyses of variance (ANOVA) were carried out for each year separately (due to differences in sampling intensity) to determine whether beetle abundance (log(x + 1)-transformed to achieve normality), richness, or Shannon’s diversity varied between treatments. Site location (= “State”) was included in the model as a blocking variable and was treated as a random effect. To provide additional information, separate analyses were performed on the following three groups: saproxylic (species directly or indirectly dependent on dead or dying wood), non-saproxylic ground beetles, and other species. These designations were based on familiarity with the taxa and information available in the literature. Voucher specimens have been deposited in the Louisiana State Arthropod Museum, Baton Rouge, Louisiana.

3. Results

The final dataset consisted of 5172 beetle specimens belonging to 55 families and 378 taxa identified to genus or species (all of which are hereafter referred to as “species”, see Appendix). Of these, 47 (13%) were non-saproxylic ground beetles, 182 (48%) were saproxylic, and 149 (39%) were other species (Appendix). Texas traps yielded the most beetles in terms of both individuals (~44% of the total) and species (~60%) whereas Louisiana traps yielded the fewest (see Appendix; note that North Carolina traps were only sampled in 1998). State (block) was found to be a significant source of variation in most analyses of overall beetle richness, abundance and diversity in 1998 and 1999 but less so in 2002 (Table 1). The same was true when saproxylic species, ground beetles (which are non-saproxylic), and other species were analyzed separately (Table 1)

With respect to treatment differences, 2998 (58%) individuals and 302 (80%) species were collected from the dead wood plots compared to 2174 (42%) individuals and 261 (69%) species from the reference plots. Furthermore, 115 and 74 species were collected from the dead wood and reference plots, respectively. Overall, beetles were consistently more species rich, abundant, and diverse in the dead wood plots than in the reference plots during the entire study (Figure 1). Although these differences were greatest in 1998, significant differences were detected only in 1999 and only for diversity (Table 1). Similarly, only in 1999 were significant differences detected between treatments for saproxylic beetles, with them being more rich and diverse in dead wood plots than in references plots. No significant differences were detected in 2002 and non-saproxylic ground beetles and other beetle species, when analyzed separately, did not vary between treatments in any year (Table 1).

The large differences in mean beetle richness, abundance and diversity between treatments in 1998 (Figure 1) were not statistically significant due to strong interactions between state and treatment (Table 1). Most notably, the results from North Carolina were not consistent with those from the other states (Figure 2, Figure 3, Figure 4). After excluding North Carolina from the dataset, the differences in overall beetle richness (F1,2 = 19.4, p < 0.05), saproxylic beetle richness (F1,2 = 23.8, p < 0.05) and saproxylic beetle diversity (F1,2 = 20.1, p < 0.05) observed in 1998 were found to be significant.

Table 1. Results from analyses of variance for all species combined, saproxylic species, non-saproxylic ground beetles and other species. Note that data from North Carolina are limited to 1998. Asterisks denote significant p-values: * < 0.05, ** < 0.01, *** < 0.001 based on analyses of variance.

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Table 1. Results from analyses of variance for all species combined, saproxylic species, non-saproxylic ground beetles and other species. Note that data from North Carolina are limited to 1998. Asterisks denote significant p-values: * < 0.05, ** < 0.01, *** < 0.001 based on analyses of variance.
Parameter1998 ( n = 12)1999 ( n = 9)2002 ( n = 9)
All speciesF3,16 = 7.5**F1,3 = 3.4F3,16 = 3.3*F2,12 = 16.4***F1,2 = 5.9F2,12 = 0.1F2,12 = 8.1**F1,2 = 1.1F2,12 = 2.3
Saproxylic speciesF3,16 = 0.9F1,3 = 6.7F3,16 = 4.5*F2,12 = 18.0***F1,2 = 45.1*F2,12 = 0.1F2,12 = 8.9**F1,2 = 1.2F2,12 = 1.3
Ground beetlesF3,16 = 14.3***F1,3 = 0.4F3,16 = 0.5F2,12 = 8.7**F1,2 = 0.1F2,12 = 0.5F2,12 = 1.5F1,2 = 1.6F2,12 = 1.8
Other speciesF3,16 = 28.6***F1,3 = 1.9F3,16 = 1.1F2,12 = 10.2**F1,2 = 5.8F2,12 = 0.1F2,12 = 3.9F1,2 = 0.6F2,12 = 1.7
All speciesF3,16 = 13.0***F1,3 = 8.1F3,16 = 2.2F2,12 = 18.5***F1,2 = 0.4F2,12 = 0.4F2,12 = 2.2F1,2 = 1.1F2,12 = 1.9
Saproxylic speciesF3,16 = 4.0*F1,3 = 8.3F3,16 = 3.6*F2,12 = 18.7***F1,2 = 1.9F2,12 = 0.9F2,12 = 6.3*F1,2 = 1.7F2,12 = 1.3
Ground beetlesF3,16 = 18.5***F1,3 = 0.4F3,16 = 0.7F2,12 = 2.7F1,2 = 0.7F2,12 = 0.8F2,12 = 2.0F1,2 = 2.4F2,12 = 0.9
Other speciesF3,16 = 11.1***F1,3 = 0.0F3,16 = 0.4F2,12 = 13.3***F1,2 = 0.1F2,12 = 0.2F2,12 = 1.7F1,2 = 0.3F2,12 = 2.5
All speciesF3,16 = 5.1*F1,3 = 2.3F3,16 = 4.7*F2,12 = 9.9**F1,2 = 147.8**F2,12 = 0.0F2,12 = 0.8F1,2 = 1.3F2,12 = 1.0
Saproxylic speciesF3,16 = 1.5F1,3 = 4.2F3,16 = 4.1*F2,12 = 1.8F1,2 = 62.4*F2,12 = 0.1F2,12 = 1.8F1,2 = 0.8F2,12 = 0.6
Ground beetlesF3,16 = 7.5**F1,3 = 0.1F3,16 = 0.4F2,12 = 16.3***F1,2 = 3.0F2,12 = 0.2F2,12 = 1.0F1,2 = 1.3F2,12 = 2.0
Other speciesF3,16 = 10.2***F1,3 = 0.2F3,16 = 1.4F2,12 = 10.3**F1,2 = 9.9F2,12 = 0.2F2,12 = 0.9F1,2 = 0.9F2,12 = 0.9
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Figure 1. Mean ± SE (n = 12 for 1998 and 9 for 1999 and 2002) species richness, abundance, and Shannon’s diversity of beetles collected in plots in which dead wood was added or not (open and closed circles, respectively). Results for all beetle species combined are given in the left-most column followed by saproxylic species only, ground beetles only, and other species. Asterisks denote significant p-values: * < 0.05, ** < 0.01, *** < 0.001 based on analyses of variance.

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Figure 1. Mean ± SE (n = 12 for 1998 and 9 for 1999 and 2002) species richness, abundance, and Shannon’s diversity of beetles collected in plots in which dead wood was added or not (open and closed circles, respectively). Results for all beetle species combined are given in the left-most column followed by saproxylic species only, ground beetles only, and other species. Asterisks denote significant p-values: * < 0.05, ** < 0.01, *** < 0.001 based on analyses of variance.
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Figure 2. Mean ± SE (n = 3) saproxylic (top) and total (bottom) beetle species richness in reference and dead wood plots (closed and open circles, respectively) by state in 1998.

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Figure 2. Mean ± SE (n = 3) saproxylic (top) and total (bottom) beetle species richness in reference and dead wood plots (closed and open circles, respectively) by state in 1998.
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Figure 3. Mean ± SE (n = 3) saproxylic (top) and total (bottom) beetle diversity (Shannon’s diversity) in reference and dead wood plots (closed and open circles, respectively) by state in 1998.

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Figure 3. Mean ± SE (n = 3) saproxylic (top) and total (bottom) beetle diversity (Shannon’s diversity) in reference and dead wood plots (closed and open circles, respectively) by state in 1998.
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Figure 4. Mean ± SE (n = 3) saproxylic beetle abundance in reference and dead wood plots (closed and open circles, respectively) by state in 1998.

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Figure 4. Mean ± SE (n = 3) saproxylic beetle abundance in reference and dead wood plots (closed and open circles, respectively) by state in 1998.
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4. Discussion

Despite conflicting results from North Carolina which we are unable to explain based on this study, beetles generally responded positively to the dead wood manipulations made in this study, particularly during the first year. Saproxylic species were more abundant, whereas non-saproxylic ground beetles and other species did not noticeably differ. The increased abundance of saproxylic species is not surprising as these organisms, by definition, are dependent on dying and dead wood [4]. Many were no doubt captured as they attempted to colonize the fresh wood added to the plots. It is noteworthy that the degree to which saproxylic species were more numerous in the dead wood plots declined after the first year. This is largely due to the fact that more species are associated with the phloem of freshly-killed loblolly pine wood than later decay stages [29]. Once the phloem is gone, which happens quickly, so too are these early colonists. Ulyshen and Hanula [29] provided a list of beetle species found to be specifically associated with freshly-killed loblolly pine in South Carolina. Of those, the following genera were collected only during 1998 (the year the wood was added to the plots) in the current study: the cerambycids Acanthocinus and Monochamus; the zopherids Colydium and Lasconotus; the curculionids Dendroctonus, Gnathotrichus, Ips, Orthotomicus, and Myoplatypus; the tenebrionid Corticeus, the staphylinid Myrmecocephalus, and the histerid Platysoma (results not shown). A wide variety of other species colonize loblolly pine after the phloem stage but not in such great numbers [29]. Furthermore, many of these species are likely to re-infest the same piece of wood until it no longer provides a suitable substrate [30], thereby reducing the likelihood of being captured by pitfall traps. While the ethanol used in the trap collection jars may have attracted insects, these effects would seem to have been equal between controls and treatments, and the added wood apparently overcame any confounding effects of using this preservative.

That ground beetles were not more strongly affected by the dead wood additions is somewhat surprising considering that many previous studies have shown this group to be positively associated with dead wood [26,27,31,32,33,34,35]. Our results, therefore, provide little support for the idea that epigeic predators are more likely than many non-saproxylic taxa to benefit from dead wood due to its positive effect on prey abundance [27]. It is possible however that the debris in the dead wood plots impeded carabid movement enough to reduce their capture rate, thereby masking a stronger beneficial effect. Furthermore, it appears that ground beetles were becoming increasingly numerous in the dead wood plots relative to the reference plots over the course of this study (see Figure 1). Although the differences were not statistically significant, ground beetle richness and diversity had non-overlapping standard errors in 2002. These findings suggest that the benefits of dead wood to these taxa may be somewhat delayed. The results from other studies are not consistent with this conclusion, however. Within the first year after sites were clearcut in Sweden, for example, Nittérus and Gunnarsson [34] collected significantly more ground beetles in pitfall traps placed under piles of slash compared to those placed out in the open. Perhaps dead wood represents a more important source of shelter from sunlight and other desiccating conditions in clearcuts than in forested plots such as those used in the current study.

5. Conclusions

While some degree of caution is advised in interpreting the statistical tests we report here (due to the relatively close proximity of the sites to one another), our results are largely consistent with those of Hanula et al. [26] and Ulyshen and Hanula [27]. These studies suggest that ground-dwelling arthropods (saproxylic species notwithstanding) are little-affected by manipulations of dead wood in loblolly pine forests. This conclusion, however, is in conflict with evidence that many ground-dwelling arthropod taxa are more numerous immediately next to loblolly pine logs than short distances away from them [15]. As suggested by Ulyshen and Hanula [27], this discrepancy may indicate that dead wood has a stronger effect on how ground-dwelling arthropods are spatially distributed (i.e., causing them to become more clumped) than on their abundance or species richness. Studies aimed at addressing this question would be of particular interest. More research is also needed to determine how manipulations of dead wood at larger scales and over longer time periods affect these organisms.


Table A1. Total beetle abundance by species, state, and treatment (reference/wood addition). Group designations are as follows: S = saproxylic (i.e., directly or indirectly dependent on dead wood at some life stage); P = predator (i.e., non-saproxylic ground beetles); O = other. Note that data from North Carolina are limited to 1998.

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Table A1. Total beetle abundance by species, state, and treatment (reference/wood addition). Group designations are as follows: S = saproxylic (i.e., directly or indirectly dependent on dead wood at some life stage); P = predator (i.e., non-saproxylic ground beetles); O = other. Note that data from North Carolina are limited to 1998.
SpeciesGroupLouisianaMississippiNorth CarolinaTexasTotal
Zonantes fasciatus (Melsheimer)S0/00/01/00/01/0
Zonantes signatus (Haldeman)S0/00/10/00/00/1
Zonantes subfasciatus (LeConte)S0/01/00/00/01/0
Zonantes sp.S0/00/10/00/00/1
Necrophilus pettitii HornO0/05/20/00/05/2
Cryptoramorphus sp.S0/00/00/00/10/1
Euvrilletta peltata (Harris)S0/00/00/00/10/1
Ptinus sp.S0/01/00/01/22/2
Tomoderus sp.O0/02/00/00/02/0
Vacusus sp.O0/01/00/00/01/0
Pterocolus ovatus (Fabricius)O0/00/00/00/10/1
Diplocoelus rudis (LeConte)S8/712/278/710/838/49
Lichenophanes bicornis (Weber)S0/00/00/00/10/1
Melalgus plicatus (LeConte)S0/00/00/00/10/1
Stephanopachys sp.S0/00/00/00/10/1
Xylobiops basilaris (Say)S0/31/20/00/31/8
Bothrideres cryptus StephanS0/00/00/00/10/1
Sayapion segnipes (Say)O0/01/00/00/01/0
Buprestis lineata FabriciusS0/10/00/00/00/1
Chalcophora virginiensis (Drury)S0/10/10/00/10/3
Rhagonycha sp.O3/00/10/00/13/2
Acupalpus rectangulus ChaudoirP0/00/00/01/01/0
Agonum punctiforme SayP0/00/00/06/156/15
Amara impuncticollis (Say)P0/00/00/00/10/1
Anisodactylus haplomus ChaudoirP3/02/00/01/06/0
Apenes lucidulus ChaudoirP0/00/10/00/00/1
Apenes sinuatus (Say)P0/00/00/10/30/4
Brachinus alternans DejeanP0/00/00/10/00/1
Brachinus americanus (LeConte)P0/00/00/00/10/1
Calathus opaculus LeConteP0/00/01/034/4335/43
Calosoma scrutator (Fabricius)P0/00/00/02/22/2
Carabus goryi DejeanP0/00/01/00/01/0
Chlaenius amoenus (Dejean)P3/22/20/00/05/4
Chlaenius emarginatus SayP0/00/02/10/02/1
Chlaenius erythropus GermarP0/00/00/01/01/0
Clivina ferrea LeConteP0/00/10/00/00/1
Clivina pallida SayP0/00/10/00/00/1
Coptodera aerata DejeanP0/21/21/21/23/8
Cyclotrachelus alabamae (Van Dyke)P46/390/20/023/2469/65
Cyclotrachelus convivus (LeConte)P0/023/300/00/023/30
Cyclotrachelus laevipennis (LeConte)P1/01/10/00/02/1
Cyclotrachelus seximpressus (LeConte)P1/10/00/00/01/1
Cyclotrachelus sigillatus (Say)P0/00/030/310/030/31
Cyclotrachelus spoliatus (Newman)P0/00/019/140/019/14
Cyclotrachelus texensis (Freitag)P6/50/00/042/5948/64
Dicaelus crenatus LeConteP0/00/062/570/062/57
Dicaelus elongatus BonelliP0/15/93/67/515/21
Dicaelus furvus DejeanP1/01/43/10/05/5
Dicaelus purpuratus BonelliP0/00/03/10/03/1
Elaphropus granarius (Dejean)P0/20/00/00/00/2
Galerita bicolor DruryP0/04/1057/820/061/92
Harpalus rufipes DegeerP0/00/00/01/01/0
Harpalus sp.P0/00/00/01/01/0
Helluomorphoides nigripennis DejeanP5/638/274/20/047/35
Helluomorphoides praestus bicolor (Larochelle and Lariviere)P0/00/10/00/00/1
Lebia ornata SayP0/00/00/10/00/1
Megacephala virginica (Linneaus)P0/10/00/00/00/1
Mioptachys flavicauda (Say)S1/10/00/00/41/5
Notiophilus novemstriatus LeConteP0/06/60/02/28/8
Panagaeus fasciatus SayP0/00/00/01/01/0
Pasimachus sublaevis (Beauvois)P0/00/02/10/02/1
Pterostichus permundus (Say)P0/00/00/04/24/2
Rhadine larvalis LeConteP0/00/20/00/00/2
Scaphinotus cavicollis (LeConte)P0/00/00/04/54/5
Scaphinotus liebecki Van DykeP0/00/00/01/01/0
Selenophorus ellipticus DejeanP1/00/00/00/01/0
Selenophorus opalinus (LeConte)P1/11/00/01/03/1
Selenophorus sp.P0/00/00/00/10/1
Sphaeroderus stenostomus (Weber)P0/00/02/00/02/0
Tachyta nana (Gyllenhal)S0/10/00/00/00/1
Acanthocinus obsoletus (Olivier)S0/100/10/70/20/20
Anelaphus pumilus (Newman)S0/00/00/00/10/1
Arhopalus rusticus nubilus (LeConte)S0/10/00/00/00/1
Astylopsis perplexa (Haldeman)S0/10/00/00/00/1
Curius dentatus NewmanS0/01/00/00/11/1
Distenia undata (Fabricius)S0/00/00/01/01/0
Eburia quadrigeminata (Say)S0/00/00/01/01/0
Elaphidion mucronatum (Say)S1/00/10/02/53/6
Enaphalodes atomarius (Drury)S1/00/00/00/01/0
Eupogonius tomentosus (Haldeman)S0/01/00/02/03/0
Graphisurus fasciatus (DeGeer)S0/10/01/53/44/10
Knulliana cincta (Drury)S0/00/00/01/51/5
Leptostylus transversus (Gyllenhal)S2/36/40/012/2820/35
Monochamus carolinensis (Olivier)S0/00/00/10/00/1
Neoclytus acuminatus (Fabricius)S0/12/11/010/013/2
Obrium maculatum (Olivier)S0/10/00/09/79/8
Orthosoma brunneum (Forster)S0/01/00/02/03/0
Prionus pocularis DalmanS0/21/130/02/33/18
Sternidius alpha (Say)S0/00/03/10/03/1
Styloleptus biustus (LeConte)S0/01/20/13/64/9
Typocerus lunulatus (Swederus)S0/00/00/01/01/0
Typocerus velutinus (Olivier)S0/01/00/00/01/0
Typocerus zebra (Olivier)S0/02/21/00/03/2
Xylotrechus colonus (Fabricius)S0/00/00/03/43/4
Xylotrechus s. sagittatus (Germar)S0/00/10/00/00/1
Philothermus glabriculus LeConteS0/00/10/00/10/2
Capraita circumdata (Randall)O0/00/30/00/10/4
Capraita obsidiana (Fabricius)O0/00/00/00/10/1
Capraita suturalis (Fabricius)O0/00/20/00/00/2
Capraita thyamoides (Crotch)O0/00/00/03/13/1
Capraita sp.O0/00/02/00/32/3
Graphops curtipennis (Melsheimer)O3/10/00/00/03/1
Graphops floridanus BlakeO2/10/00/00/02/1
Metachroma pellucidum CrotchO0/00/20/00/00/2
Orthaltica copalina (Fabricius)O0/01/00/00/01/0
Paria sp.O0/40/00/012/312/7
Rhabdopterus sp.O0/03/02/10/05/1
Cis sp.S0/00/00/01/11/1
Cymatodera wolcotti BarrS1/00/00/00/01/0
Neorthopleura thoracica (Say)S0/10/20/01/21/5
Priocera castanea (Newman)S0/50/10/00/00/6
Psyllobora vigintimaculata (Say)O0/00/01/00/01/0
Arthrolips fasciata (Erichson)S0/00/00/00/30/3
Clypastracea sp.S0/00/00/00/10/1
Cryptophagus sp.S1/00/00/00/11/1
Acalles clavatus (Say)S0/13/193/31/27/25
Ambrosiodmus rubricolllis (Eichhoff)S1/15/811/133/520/27
Apteromechus ferratus (Say)S0/00/41/113/2714/32
Coccotrypes distinctus (Motschulsky)S0/10/00/00/00/1
Conotrachelus posticatus BohemanO0/00/01/0103/68104/68
Cophes fallax (LeConte)S0/00/00/01/31/3
Corthylus punctatissimus (Zimmermann)S0/01/00/10/01/1
Cossonus corticola SayS0/131/60/05/416/60
Cryptorhynchus tristis LeConteO1/22/30/01/04/5
Cyrtepistomus castaneus (Roelofs)O0/01/00/00/01/0
Dendroctonus terebrans (Olivier)S0/00/00/00/60/6
Dryophthorus americanus BedelS0/22/310/04/316/8
Dryoxylon onoharaensis (Murayama)S0/01/40/00/01/4
Euplatypus compositus (Say)S4/70/20/01/15/10
Euwallacea validus (Eichhoff)S0/00/10/00/00/1
Gnathotrichus materiarius (Fitch)S0/00/00/30/10/4
Hylastes porculus ErichsonS0/20/00/10/10/4
Hylastes salebrosus EichhoffS1/20/00/00/21/4
Hylastes tenuis EichhoffS1/21/96/91/99/29
Hylobius pales (Herbst)S2/88/1211/303/1224/62
Hypothenemus sp.S0/00/01/10/11/2
Ips avulsus (Eichhoff)S0/21/10/01/52/8
Ips calligraphus (Germar)S0/10/10/00/20/4
Ips grandicollis (Eichhoff)S0/00/00/00/40/4
Lissorhoptrus oryzophilus KuschelO0/10/00/00/00/1
Lissorhoptrus simplex (Say)O1/00/00/00/01/0
Monarthrum fasciatum (Say)S0/00/00/00/10/1
Monarthrum mali (Fitch)S0/00/00/02/42/4
Myoplatypus flavicornis (Fabricius)S0/150/30/250/190/62
Notaris puncticollis (LeConte)O0/01/00/00/01/0
Orthotomicus caelatus (Eichhoff)S0/387/1011/5225/5943/159
Pachylobius picivorus (Germar)S5/3010/384/1325/4044/121
Pityophthorus confusus BlandfordS0/10/00/00/00/1
Sphenophorus sp.O0/00/00/01/01/0
Stethobaris sp.O1/10/00/00/01/1
Xyleborinus saxeseni (Ratzeburg)S14/1022/129/2117/98162/122
Xyleborus affinis EichhoffS4/3628/5127/1813/6072/165
Xyleborus ferrugineus (Fabricius)S3/336/1623/1039/5871/117
Xyleborus pubescens ZimmermannS1/30/01/11/53/9
Xyleborus xylographus (Say)S0/00/01/00/11/1
Xylosandrus compactus (Eichhoff)S0/01/122/301/224/33
Xylosandrus crassiusculus (Motschulsky)S1/49/179/90/519/35
Xylosandrus germanus (Blandford)S1/04/70/00/05/7
Copelatus glyphicus (Say)O5/10/01/11/17/3
Alaus myops (Fabricius)S0/10/10/00/00/2
Blauta cribraria (Germar)S0/30/10/00/00/4
Conoderus amplicollis (Gyllenhal)S0/10/00/00/00/1
Dicrepidius sp.S0/00/00/00/10/1
Glyphonyx bimarginatus SchaefferS0/00/00/01/01/0
Glyphonyx ferruginosus SchaefferS0/00/00/01/11/1
Glyphonyx sp.S0/00/00/10/00/1
Lacon discoideus (Weber)S0/00/10/00/00/1
Lacon impressicollis (Say)S0/00/00/00/10/1
Limonius quercinus SayS0/00/01/00/01/0
Megapenthes rufilabris (Germar)S1/00/10/00/11/2
Megapenthes sp.S0/00/00/10/00/1
Melanotus corticinus (Say)S0/00/10/00/00/1
Melanotus ignobilis MelsheimerS1/20/00/00/01/2
Melanotus insipiens (Say)S0/00/00/01/01/0
Melanotus piceatus BlatchleyS0/00/02/00/02/0
Melanotus pilosus BlatchleyS1/10/00/00/01/1
Melanotus similis groupS0/01/10/10/01/2
Melanotus testaceus (Melsheimer)S0/00/00/01/01/0
Melanotus sp.S0/02/20/10/12/4
Mulsanteus carolinensis (Schaeffer)S0/00/00/013/213/2
Aphorista vittata (Fabricius)S1/22/30/04/77/12
Danae testacea (Ziegler)S0/12/37/34/313/10
Epipocus punctatus LeConteS1/10/00/00/01/1
Lycoperdina ferruginea LeConteS0/04/21/08/1213/14
Mycetina perpulchra (Newman)S0/00/10/00/00/1
Cryptophilus integer (Heer)S0/00/10/00/00/1
Triplax festiva LacordaireS0/00/00/01/01/0
Tritoma affinis LacordaireS2/311/30/08/1121/17
Tritoma angulata SayS0/01/00/00/01/0
Tritoma atriventris LeConteS0/00/00/01/31/3
Tritoma biguttata affinis LacordaireS1/24/40/04/39/9
Tritoma humeralis FabriciusS0/00/00/30/00/3
Eucinetus strigosus LeConteS1/44/70/00/05/11
Dromaeolus cylindricollis (Say)S5/01/20/01/27/4
Dromaeolus striatus (LeConte)S1/12/20/00/03/3
Microrhagus triangularis (Say)S0/00/00/02/12/1
Bolboceras thoracicornis (Wallis)O0/00/01/00/01/0
Bolbocerosoma farctum (Fabricius)O0/01/10/00/01/1
Geotrupes blackburnii (Fabricius)O1/00/00/01/32/3
Geotrupes opacus HaldemanO0/00/00/16/06/1
Odonteus sp.O0/00/01/00/01/0
Eblisia carolina (Paykull)S0/00/00/00/10/1
Paromalus seminulum ErichsonS0/11/00/00/11/2
Platysoma coarctatum LeConteS0/00/10/00/20/3
Cercyon occallatus (Say)O0/01/09/40/010/4
Cercyon pubescens LeConteO1/00/00/03/04/0
Cymbiodyta chamberlaini SmetanaO1/00/00/00/11/1
Cryptolestes punctatus (LeConte)S0/10/00/00/00/1
Cryptolestes sp.S0/00/00/00/10/1
Laemophloeus biguttatus SayS0/00/10/00/00/1
Placonotus modestus (Say)S0/00/20/04/24/4
Ellychnia corrusca (LeConte)S0/00/01/00/01/0
Aridius sp.S0/00/00/10/00/1
Corticarina sp.S1/20/00/00/01/2
Anisotoma basalis (LeConte)O0/00/01/00/01/0
Anisotoma discolor (Melsheimer)O0/00/01/01/52/5
Anisotoma sp.O0/00/00/01/01/0
Colenis bifida PeckO0/00/02/00/02/0
Colenis impunctata LeConteO2/100/15/819/1626/35
Colenis ora PeckO0/12/10/015/2017/22
Colenis stephani PeckO0/00/00/02/12/1
Colenis sp.O0/10/00/10/00/2
Dissochaetus oblitus (LeConte)O0/00/00/10/00/1
Leiodes stephani BaranowskiO0/00/00/04/04/0
Ptomaphagus consobrinus LeConteO4/50/00/03/57/10
Ptomaphagus sp.O0/00/00/01/01/0
Dorcus parallelus (Say)S0/00/01/10/01/1
Plateros sp.S0/21/30/01/02/5
Dircaea liturata (LeConte)S0/00/00/00/10/1
Microtonus sericans LeConteS0/01/00/00/01/0
Mordella sp.O0/01/10/00/11/2
Mordellaria borealis (LeConte)O0/11/00/00/01/1
Mordellaria serval (Say)O0/01/00/01/32/3
Mordellistena pubescens (Fabricius)O0/00/10/00/00/1
Typhaea stercorea (Linnaeus)S0/21/00/017/1918/21
Amphicrossus ciliatus (Olivier)O0/00/10/00/00/1
Carpophilus antiquus MelsheimerO0/00/01/11/02/1
Carpophilus sp.O1/10/21/43/25/9
Colopterus unicolor (Say)O10/1612/3943/5530/6695/176
Epuraea helvola ErichsonO0/01/03/130/04/13
Epuraea planulata ErichsonO0/00/00/10/00/1
Epuraea rufa (Say)O0/00/07/40/07/4
Glischrochilus sanguinolentus (Olivier)O0/01/20/00/01/2
Pallodes austrinus LeschenO0/02/01/01/24/2
Pallodes pallidus BeauvoisO0/01/31/30/02/6
Prometopia sexmaculata (Say)O0/10/00/01/01/1
Stelidota coenosa ErichsonO8/72/31/123/2234/33
Stelidota geminata (Say)O1/30/17/126/314/19
Stelidota octomaculata (Say)O0/00/00/00/20/2
Stelidota sexmaculata (Say)O0/00/00/01/01/0
Stelidota strigosa SchoenherrO0/01/00/06/37/3
Oxycopis notoxoides (Fabricius)O1/00/00/00/01/0
Oxycopis sp.S0/00/00/01/01/0
Oxycopis thoracica (Fabricius)S0/00/10/00/00/1
Catogenus rufus (Fabricius)S0/10/00/00/00/1
Ptilodactyla sp.O0/00/00/02/42/4
Anomala sp.O0/00/00/00/20/2
Ataenius imbricatus MelsheimerO0/00/10/00/00/1
Ataenius insculptus HornO0/02/10/00/02/1
Ataenius sp.O0/01/00/00/01/0
Ateuchus histeroides WeberO0/00/00/02/12/1
Canthon ebenus SayO0/00/00/07/77/7
Canthon viridis (Beauvois)O0/00/10/01/11/2
Copris minutus (Drury)O0/01/00/20/01/2
Deltochilum gibbosum (Fabricius)O0/00/03/010/313/3
Digitonthophagus gazella (Fabricius)O0/01/00/00/01/0
Diplotaxis sp.O0/05/10/03/68/7
Euphoria sepulcralis (Fabricius)O0/00/00/02/72/7
Melanocanthon sp.O0/00/00/02/52/5
Onthophagus hecate (Panzer)O0/01/00/010/711/7
Onthophagus medorensis BrownO0/00/00/031/1631/16
Onthophagus pennsylvanicus HaroldO0/00/00/02/02/0
Onthophagus striatulus (Beauvois)O0/12/00/02/24/3
Onthophagus tuberculifrons HaroldO0/00/00/00/20/2
Onthophagus sp.O0/00/00/00/10/1
Parataenius simulator (Harold)O0/00/00/00/10/1
Phyllophaga forsteri (Burmeister)O0/01/20/00/01/2
Phyllophaga prunina (LeConte)O2/30/00/00/32/6
Phyllophaga prununculina (Burmeister)O0/00/10/00/00/1
Phyllophaga scitula (Horn)O0/00/00/00/10/1
Phyllophaga sp.O1/00/01/10/02/1
Phyllophaga tristis complexO0/00/00/01/01/0
Platytomus longulus (Cartwright)O1/00/00/00/01/0
Serica sp.O0/00/00/02/02/0
Scraptia sp.O0/00/01/00/01/0
Necrophila americana (Linnaeus)O0/00/00/00/20/2
Ahasverus advena (Waltl)S0/00/00/00/10/1
Ahasverus rectus (LeConte)S1/121/61/07/1410/32
Cathartosilvanus imbellis (LeConte)S0/00/10/00/10/2
Silvanoprus scuticollis (Walker)S0/00/00/00/10/1
Silvanus muticus SharpS1/20/00/02/63/8
Sphindus sp.S2/01/03/01/47/4
Achenomorphus corticinus (Gravenhorst)O1/012/110/40/113/16
Anotylus sp.O0/10/00/027/3927/40
Arthmius sp.O0/00/01/00/01/0
Astenus linearis (Erichson)O0/10/00/00/00/1
Astenus sp.O1/22/00/00/23/4
Baeocera laevis (Reitter)O0/00/00/01/01/0
Baeocera sp.O0/05/20/11/46/7
Belonuchus ephippiatus (Say)O1/00/10/01/02/1
Belonuchus rufipennis (Fabricius)S0/00/01/00/01/0
Bryoporus rufescens LeConteO0/03/10/10/03/2
Carpelimus sp.O0/10/00/00/10/2
Coproporus laevis LeConteS0/00/10/00/00/1
Coproporus ventriculus (Say)S1/00/00/00/01/0
Ctenisodes sp.O0/00/00/07/07/0
Decarthron sp.O1/00/00/00/01/0
Diochus schaumi KraatzO0/00/00/00/10/1
Echiaster sp.O0/00/10/00/10/2
Erichsonius sp.O0/10/00/00/00/1
Euconnus sp.O0/10/02/00/12/2
Hesperus baltimorensis (Gravenhorst)O0/20/02/00/02/2
Homaeotarsus sp.O0/00/00/01/01/0
Hoplandria laevicollis (Notman)O0/00/05/30/05/3
Hoplandria sp.O0/00/112/191/013/20
Ischnosoma flavicolle (LeConte)O1/01/10/01/23/3
Myrmecocephalus concinnus (Erichson)S0/20/00/00/00/2
Oxybleptes davisi (Notman)O37/428/230/00/065/27
Oxypoda sp.O0/00/00/01/01/0
Palaminus sp.S0/01/20/02/03/2
Philonthus umbrinus (Gravenhorst)O0/00/01/10/01/1
Philotermes sp.S0/09/50/00/09/5
Pinophilus sp.O1/00/00/00/01/0
Platydracus fossator (Gravenhorst)O0/00/01/02/13/1
Platydracus sp.O0/00/00/02/12/1
Quedius capucinus (Gravenhorst)O0/00/00/00/10/1
Quedius verres SmetanaO0/08/110/05/213/13
Rugilus sp.O0/03/00/02/95/9
Scaphisoma punctulatum LeConteO0/00/10/00/00/1
Scydmaenus sp.O0/00/02/00/02/0
Sepedophilus basalis (Erichson)O0/00/00/01/01/0
Sepedophilus crassus (Gravenhorst)O0/00/00/10/00/1
Sepedophilus debilis (Casey)O0/00/01/00/01/0
Sepedophilus sp.O22/182/20/113/1937/40
Stenichnus sp.O0/00/01/00/01/0
Tachinus fimbriatus GravenhorstO0/00/02/00/02/0
Thoracophorus costalis (Erichson)S0/00/00/030/2630/26
Tmesiphorus costalis LeConteO0/00/10/00/00/1
Alobates morio (Fabricius)S0/00/00/01/01/0
Alphitophagus bifasciatus (Say)S0/20/10/00/00/3
Anaedus brunneus (Ziegler)O1/00/10/00/01/1
Corticeus thoracicus (Melsheimer)S0/20/00/00/10/3
Gondwanocrypticus obsoletus (Say)S1/00/00/00/01/0
Helops cisteloides (Germar)S0/00/00/01/11/1
Helops sp.S0/00/00/01/01/0
Hymenorus sp.S3/41/20/03/57/11
Isomira sp.S0/00/00/00/20/2
Lobopoda erythrocnemis (Germar)S3/32/00/011/716/10
Opatrinus minimus (Beauvois)O0/00/00/01/11/1
Platydema micans ZimmermanS2/10/00/02/24/3
Platydema ruficolle Laporte and BrulléS0/00/10/02/12/2
Platydema ruficorne (Sturm)S0/10/10/00/00/2
Polypleurus perforatus (Germar)S0/11/00/04/25/3
Statira gagatina (Melsheimer)O0/00/10/00/00/1
Statira sp.O0/00/00/00/20/2
Uloma imberbis LeConteS0/00/02/00/12/1
Uloma punctulata LeConteS0/62/40/00/12/11
Eustrophopsis bicolor (Fabricius)S1/10/00/00/11/2
Eustrophus tomentosus SayS0/00/00/01/01/0
Aulonothroscus convergens (Horn)O0/00/40/00/00/4
Omorgus monachus HerbstO0/00/00/00/10/1
Trox spinulosus RobinsonO0/00/00/02/02/0
Trox variolatus MelsheimerO1/10/00/01/12/2
Temnoscheila virescens (Fabricius)S0/20/00/01/01/2
Bitoma quadriguttata (Say)S0/00/10/00/00/1
Colydium lineola SayS0/00/00/00/10/1
Colydium nigripenne LeConteS0/30/90/00/30/15
Endeitoma dentata (Horn)S0/11/00/00/01/1
Endeitoma sp.S0/10/00/00/00/1
Hyporhagus punctulatus ThomsonS0/00/00/00/10/1
Lasconotus pusillus LeConteS0/00/00/00/40/4
Microsicus parvulus (Guerin)S0/00/00/04/14/1
Pycnomerus haematodes (Fabricius)S0/00/00/00/10/1
Pycnomerus sulcicollis LeConteS0/42/90/00/12/14
Total Individuals (reference/wood addition) 7531016110422995172
Total Species (reference/wood addition) 140162108227378


We thank Jacob Carpenter, Erich Vallery, Rick Stagg, and cooperators in TX, NC, and MS for technical assistance. Victoria Bayless, Stephanie Gil, Jong-Seok Park, Igor Sokolov, and Alexey Tishechkin also assisted with identifications and curatorial support. We also thank Jim Hanula, Tim Schowalter and Cavell Brownie for commenting on an early version of the manuscript.

Conflict of Interest

The authors declare no conflict of interest.


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