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Ground Beetle (Coleoptera: Carabidae) Species Composition in the Southern Appalachian Mountains
Robert Browne, Sarah Maveety, Leigh Cooper, and Kathryn Riley

Southeastern Naturalist, Volume 13, Issue 2 (2014): 407–422

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Southeastern Naturalist 407 R. Browne, S. Maveety, L. Cooper, and K. Riley 22001144 SOUTHEASTERN NATURALIST 1V3o(2l.) :1430,7 N–4o2. 22 Ground Beetle (Coleoptera: Carabidae) Species Composition in the Southern Appalachian Mountains Robert Browne1,*, Sarah Maveety1, Leigh Cooper1, and Kathryn Riley1 Abstract - Using pitfall traps on 12 sites in the southern Appalachian Mountains during 2007–2008, we collected 6552 carabid beetles representing 46 species. We collected 40 species in 14 genera at 9 spruce–fir sites and 29 species in 12 genera at 3 hardwood sites. When adjusted for sampling effort via rarefaction, spruce–fir and hardwood sites did not differ in species richness. However, there were significant differences in species composition. Based on non-metric multidimensional scaling (NMS) analysis, species assemblages for spruce–fir forest were distinct from those found for hardwood forests, with the 4 northern spruce–fir forest sites clustered independently from the 5 southern spruce–fir sites. Composition by genera varied by season: Pterostichus was the dominant genus in the summer and autumn, and Sphaeroderus was the dominant genus in the winter and spring. The species captured by pitfall traps in this study differed somewhat from the species found at these sites in a previous survey made by hand-collection. However, when adjusted for sample size via rarefaction, species richness, evenness, and Fisher’s α did not differ between these samples made by different collection methods. Introduction Ground beetles (Order Coleoptera: Family Carabidae) have often been used as model organisms in ecological studies (Erwin 1996, Koivula et al. 2002) because they are a diverse group and are relatively straightforward to identify, with a wide range of morphologies, taxonomies, behaviors, and ecologies (Erwin 1996). Because the majority of carabid species are incapable of flight (at least in temperate zone forests) and are sensitive to environmental and ecological change (Niemelä et al. 2000), carabid assemblages might act as indicators of local biodiversity. Although studies have been conducted in a wide variety of ecosystems, much of what is known about carabid species assemblages comes from studies in northern hemisphere boreal forests (e.g., Butterfield et al. 1995; Desender et al. 1999; de Warnaffe and Lebrun 2004; Jukes et al. 2001; Koivula et al. 2002; Niemelä 1990; Niemelä et al. 1992a, 1993, 1996; Paquin 2008). In North America, research on carabid species diversity and community composition has primarily focused on the temperate forests of the northeast US (Jennings and Tallamy 2006, Larsen et al. 2003, Lenski 1982, Liebherr and Mahar 1979). Collections and species descriptions are available for the southern Appalachian region (Barr 1985, Carlton and Bayless 2007, Darlington 1943, Kane et al. 1990, Noonan et al. 1992), though there is little research describing community composition and factors influencing species diversity in the region (but see Ortiz and Browne 2011, Worthen and 1Department of Biology and Environmental Program, Wake Forest University, Winston- Salem, NC 27104. *Corresponding author - Manuscript Editor: Wade Worthen Southeastern Naturalist R. Browne, S. Maveety, L. Cooper, and K. Riley 2014 Vol. 13, No. 2 408 Merriman 2013). Although seasonal occurrence has been reported for some North American carabid species (Larochelle and Lariviere 2003), detailed studies for carabid species assemblages have primarily been reported for north temperate sites, e.g., Alberta (Niemelä et al. 1992b). In the southern Appalachians of Virginia, North Carolina, and Tennessee, colder temperatures and high rainfall are conducive to growth of Picea rubens Sargent (Red Spruce) and Abies fraseri Pursh (Poir) (Fraser Fir), which are the dominant trees at elevations >1370 m (White et al. 1993). These spruce–fir forests only occur on mountaintops, and represent relict ecosystems from the last glacial period (Crespi et al. 2003, Ortiz and Browne 2011, Sipe and Browne 2004). As such, spruce–fir forests are insular “sky-islands” for habitat specialists such as Glaucomys sabrinus Shaw (Northern Flying Squirrel; Arbogast et al 2005, Browne et al. 1999), Demoganathus wrighti King (Pygmy Salamander; Crespi et al. 2003), and Desmognathus organi (Northern Pygmy Salamander; Crespi et al. 2003, 2010). Curiously, carabid beetles do not seem to respond to these habitats as islands. Contrary to the predictions of the theory of island biogeography (MacArthur and Wilson 1967), Ortiz and Browne (2011) found no significant relationships between habitat size or degree of isolation and any measure of carabid diversity (species richness, Shannon-Weiner diversity, Shannon’s evenness, Fisher’s alpha, probability of interspecific encounter [PIE], dominance, or rarefaction score). Ortiz and Browne (2011) collected beetles by hand, in an active-search method. Numerous studies of arthropod communities suggest that choice of collection method influences the species detected and their relative abundances. Pitfall traps have routinely been used in temperate regions to assess carabid diversity (e.g., Dufrêne and Legendre 1997, Günter and Assmann 2004, Liu et al. 2007). Pitfall traps might reflect activity patterns more than community composition (Greenslade 1964, Gutiérrez and Menéndez 1997). However, the amount of activity may be a better index of ecological importance and competitive and predatory effects than absolute abundance alone (Lenski 1982). Pitfall trapping is not always the most appropriate sampling method for carabids. A more complete inventory of the species present in a study area is obtained by employing multiple modes of collection that target different activity types and niches (Coddington et al. 1991, Longino et al. 2002, Maveety et al. 2011). For these reasons, we decided to sample these sites again using a new method. As such, the principle objectives of this investigation were to: 1) identify, via pitfall trapping, carabid species composition for spruce–fir sites in the southern Appalachian Mountains; 2) describe how community composition varies between forest habitats (spruce–fir vs. hardwood) and between seasons; 3) and compare carabid species composition in samples from pitfall traps and in samples gathered by Ortiz and Browne (2011) using hand-collection techniques. Methods Using pitfall traps at 9 sites representing the largest spruce–fir areas in the southern Appalachians, we collected ground beetles (Coleoptera: Carabidae) (Table 1, Southeastern Naturalist 409 R. Browne, S. Maveety, L. Cooper, and K. Riley 2014 Vol. 13, No. 2 Fig. 1). For more detailed site descriptions of the sites, see White et al. (1993) and Ortiz and Browne (2011). For comparative purposes, we also collected ground beetles at 3 lower-elevation hardwood sites (Table 1) that are dominated by Quercus Figure 1. Map of spruce–fir forest study sites in the southern Appalachian Mountains. Clingmans’s Dome and Newfound Gap sites are located within the Great Smoky Mountains National Park. Shaded areas indicate spruce–fir forests (after O rtiz and Browne 2011). Table 1. Description of sites, with number of carabid beetles (n) collected at each site. Area size is not given for hardwood sites since it is orders of magnitude greater than for spruce–fir sites. Forest type: spruce–fir (SF) and hardwood (HW). Forest Latitude (N), Elevation Site Code type longitude (W) (m) Area (ha) n Clingman’s Dome, NC GSM SF 35°33.473', 83°29.690' 1606–1920 18,390 1692 Grandfather Mt., VA GF SF 36°05.958', 81°47.740' 1324 285 430 Newfound Gap, NC NF SF 35°33.335', 83°08.382' 1554 931 697 Mt. Mitchell, NC MI SF 35°46.149', 82°15.818' 2008 4337 609 Mt. Rogers, VA MR SF 36°38.122', 81°36.373' 1616 639 236 Richland Balsam/ RIPI SF 35°20.700', 82°57.937' 1874 1740 959 Mt. Pisgah, NC Roan Mt., NC RM SF 36°06.461', 82°06.594' 1859 713 186 Water Rock Knob, NC WR SF 35°27.555', 83°08.382' 1798 537 514 Whitetop Mt./ WTEG SF 36°38.310', 81°36.373' 1646 138 185 Elk Garden, VA Bradley Fork Trail NC BF HW 35°33.182', 83°18.702' 675 - 297 Doughton Park, NC DP HW 36°25.555', 81°09.105' 975 - 532 Kephart Trail NC KEP HW 35°35.327', 83°21.900' 886 - 215 Southeastern Naturalist R. Browne, S. Maveety, L. Cooper, and K. Riley 2014 Vol. 13, No. 2 410 (oak) and Fagus (beech). Carabid beetles had previously been collected by hand at 12 sites (Ortiz and Browne 2011); 11 of the sites we sampled for this study were also sampled in the previous study. Due to road closure, we did not sample in the present study the Balsam (BA) site that Ortiz and Browne (2011) sampled. Newfound Gap (NF), located at approximately the same elevation within Great Smoky Mountains National Park but 20 km to the east, was substituted for BA in this study. We believe, based on forest age, elevation, rain fall, and tree species composition, that these two spruce–fir sites are approximately ecologically equivalent. The maximum distance is 206 km between any two of the spruce–fir study sites, and 224 km between any two of the hardwood study sites. We constructed each pitfall trap by embedding a 16-ounce plastic cup in the soil with the rim flush with the ground. We embedded 12 cups, arranged along an approximately 20-m line, at each site. We half-filled the cups with non-toxic antifreeze (propylene glycol), which kills and preserves arthropods. A flexible foam cover (12 x 12 cm), raised approximately 1 cm above each cup (via nails partially pushed into the soil) prevented excess rainwater and debris from entering the cup. Each month from June 2007 through May 2008, we drained the contents of each cup through a fine-mesh strainer and removed adult carabid beetles, which we preserved in 95% ethanol, identified to species, and stored at Wake Forest University. We used several indices to measure carabid beetle diversity for each site (for detailed descriptions of indices, see Magurran 2004): raw number of species, number of genera, Fisher’s alpha, and Shannon’s evenness (E). We adjusted species numbers for sampling effort using rarefaction curves. Randomized rarefaction curves without replacement were produced for each site with EstimateS 8.20 using the Mao Tau richness index (Colwell 2005). The steepness of the slope of the curve indicates the rate at which more species can be expected to be added to the sample with more collection effort. When a curve is at asymptote, it implies that all species present at that location have been collected. We analyzed species composition via non-metric multidimensional scaling (NMDS), using PC-ORD Version 6 ( NMDS was calculated based on Sørensen’s (Bray-Curtis) index, using 500 runs with real data and 100 iterations; we analyzed three dimensions and reported in the results the two dimensions with greatest r2. We also analyzed species composition for seasonal effects. We defined seasons as: summer (June–August), autumn (September–November), winter (December–February), and spring (March–May). We compared data from this pitfall trap study to previously reported data for carabids based on hand collections (Ortiz and Browne 2011). The same sites were used in both studies, with the exception of the substitution of the NF site for the BA site. Results Collections from pitfall traps We collected a total of 6552 individuals representing 46 species in 14 genera: 5508 individuals representing 40 species in 14 genera from 9 spruce–fir sites (Table 2), and 1044 carabid beetles representing 29 species in 12 genera from 3 Southeastern Naturalist 411 R. Browne, S. Maveety, L. Cooper, and K. Riley 2014 Vol. 13, No. 2 Table 2. Species composition for 9 spruce–fir sites and 3 hardwoo d sites in the southern Appalachians. See text for site abbreviations. Spruce–fir Hardwood Species GSM GF NF MI MR RIPI RM WR WTEG Total BF DP KEP Total Agonum spp. A 0 1 1 Agonum spp. B 1 1 0 Carabus goryi Dejean 10 13 1 274 22 133 8 21 23 505 139 227 2 368 Carabus serratus Say 6 6 0 Chlaenius amoenus Dejean 1 1 0 Chlaenius emarginatus Say 1 1 1 3 1 1 Cyclotrachelus laevipennis LeConte 3 1 4 1 3 4 Cyclotrachelus sigillatus Say 1 1 4 3 51 58 Dicaelus ambiguus LaFerte-Senectere 2 2 0 Dicaelus polinotes Dejean 0 1 1 Dicaelus politus Dejean 0 2 2 Dicaelus teter Bonelli 8 6 1 1 9 25 17 10 19 46 Galerita lecontei lecontei Dejean 2 2 0 Gastrellarius blanchardi Horn 2 2 1 1 Gastrellarius honestus Say 12 2 3 20 2 3 3 45 1 1 8 10 Gastrellarius unicarum Say 1 1 0 Harpalus herbivagus Say 1 1 2 0 Harpalus pensylvanicus De Geer 1 1 2 0 Harpalus spadiceus Dejean 13 5 1 4 3 2 28 2 11 13 Maronetus debilis LeConte 14 1 5 39 1 5 65 0 Myas coracinus Say 0 1 1 Myas cyanescens Dejean 3 3 0 Platynus angustatus Dejean 6 1 2 76 4 1 90 2 2 Platynus cincticollis Say 1 1 2 0 Southeastern Naturalist R. Browne, S. Maveety, L. Cooper, and K. Riley 2014 Vol. 13, No. 2 412 Table 2, continued. Spruce–fir Hardwood Species GSM GF NF MI MR RIPI RM WR WTEG Total BF DP KEP Total Platynus decentis Say 0 1 1 Platynus trifoveolatus Beutenmuller 1 1 0 Pterostichus acutipes Barr 0 7 7 Pterostichus adoxus Say 176 67 22 22 5 103 6 75 30 506 2 19 2 23 Pterostichus coracinus Newman 62 2 23 20 53 160 0 Pterostichus lachrymosus Newman 506 38 311 17 7 404 86 1369 70 70 Pterostichus moestus Say 4 1 5 4 13 17 Pterostichus mutus Say 1 3 1 5 0 Pterostichus palmi Schaeffer 2 2 3 1 8 0 Pterostichus relictus Newman 1 1 1 1 Pterostichus rostratus Newman 98 47 9 83 47 114 72 30 1 501 11 4 15 Pterostichus tristis Dejean 73 18 11 161 31 8 8 310 10 1 11 Scaphinoutus andrewsii Valentine 6 5 3 3 3 20 0 Scaphinotus elevatus Fabricius 76 9 101 186 83 2 85 Scaphinotus guyotii LeConte 1 2 2 1 17 23 1 1 Scaphinotus tricarnitus Casey 93 43 3 31 1 30 201 1 1 1 3 Scaphinotus viduus Dejean 45 2 3 3 7 2 62 1 1 Scaphinotus violaceous LeConte 2 25 2 2 2 3 2 38 3 1 4 Sphaeroderus bicarinatus LeConte 271 2 105 1 32 1 87 2 501 69 11 99 179 Sphaeroderus canadensis Chaudoir 310 21 17 5 2 45 24 32 9 465 11 10 12 33 Sphaeroderus schaumi Chaudoir 1 13 1 15 0 Sphaeroderus stenostomus Dejean 53 157 4 2 31 4 44 6 40 341 34 50 1 85 Total 1692 430 697 609 236 959 186 514 185 5508 297 532 215 1044 Southeastern Naturalist 413 R. Browne, S. Maveety, L. Cooper, and K. Riley 2014 Vol. 13, No. 2 hardwood sites. For all statistical tests, we corrected abundance for trap nights and adjusted for sampling effort based on n = 185 at all sites. Four species were present at all 12 sites: Carabus goryi, Pterostichus adoxus, Sphaeroderus canadensis, and Sphaeroderus stenostomus. These four species can be considered elevation/foresttype generalists. In addition to these four generalist species, Pterostichus rostratus was found at all spruce–fir sites, and Cyclotrachelus sigillatus, Dicaelus teter, Gastrellarius honestus, and Scaphinotus tricarnitus occured at all hardwood sites. When comparing indices for spruce–fir sites versus hardwood sites (Table 3), Shannon’s evenness was higher for spruce–fir sites than hardwood sites (single factor ANOVA: P = 0.014). Richness adjusted by rarefaction (Sadj) did not differ significantly between spruce–fir and hardwood sites. There was no difference in occurrence of rare species between spruce–fir and hardwood forests. For spruce–fir forests, 27/40 species (68%) were rare (defined as <1% of total abundance); for hardwood forests, 16/29 (55%) species were rare. For genera (Fig. 2), a higher proportion of Pterostichus and Maronetus were found in spruce–fir forests than in hardwood forests (repeated measures ANOVA: P < 0.01 and P < 0.002, respectively). Conversely, Carabus and Dicaelus were more common in hardwood forests than spruce–fir forests (repeated measures ANOVA: P < 0.001 and P < 0.0001, respectively). For carabid beetle taxa assemblages as a whole, spruce–fir and hardwood forests differed significantly (χ2 = 58.67, P < 0.001). Six species were significantly more abundant at spruce–fir sites than hardwood sites (Table 2): Pterostichus adoxus, Pterostichus coracinus, Pterostichus rostratus, Scaphinotus andrewsii, Scaphinotus tricarinitus, and Maronetus debilis Table 3. Diversity indices for carabid beetle assemblages. Upper panel is for spruce–fir sites; Lower panel is for hardwood sites. S = number of species, E = Shannon’s evenness, α = Fisher’s alpha, Sadj = species number based on rarefaction. See Table 1 for site codes. Site n S E α Sadj Spruce–fir sites GSM 1698 24 0.66 3.96 12.5 GF 430 20 0.71 4.34 12.5 MI 606 17 0.56 3.25 9.7 MR 236 18 0.71 4.53 14.0 NF 697 23 0.61 4.57 12.7 RIPI 959 18 0.66 3.14 11.5 RM 186 13 0.68 3.18 10.5 WR 514 18 0.78 3.63 13.5 WTEG 166 17 0.72 4.74 14.4 Mean 610.2 18.70 0.68 3.93 12.37 S.E. 161.2 1.10 0.02 0.22 0.52 Hardwood sites BF 297 16 0.59 3.62 11.5 DP 532 22 0.62 4.62 13.2 KEP 215 17 0.60 4.32 11.5 Mean 348 18.30 0.62 4.19 12.07 S.E. 95 1.86 0.01 0.30 0.57 Southeastern Naturalist R. Browne, S. Maveety, L. Cooper, and K. Riley 2014 Vol. 13, No. 2 414 (single factor ANOVA: P < 0.05 in all cases). Only Cyclotrachelus sigillatus was significantly more common in hardwood sites than spruce–fir sites (single factor ANOVA: P < 0.01). For NMS analysis (Fig. 3), Axes 2 and 3 had the largest r values, although Axes 1 and 2 were significantly correlated (F = 5.7569, P = 0.037, r = 0.605). Spruce–fir forest sites and hardwood forest sites clustered independently (Fig. 3). Figure 3. Nonmetric multidimensional scaling (NMS) based on Sorenson’s similarity index derived from carabid species presence/absence from our pitfall-trap collections. Figure 2. Composition of Carabidae genera obtained from traps for combined spruce–fir sites and combined hardwood sites. Genera consisting of <1.20% of collected individuals are not shown. Southeastern Naturalist 415 R. Browne, S. Maveety, L. Cooper, and K. Riley 2014 Vol. 13, No. 2 Spruce–fir forests are aligned on a northeast-to-southwest axis, with the four northern-most sites (MR, WTEG, RM, and GF) clustering in the lower left-hand portion of the quadrant, and the remaining more southern sites clustering in the upper right of the quadrant. The correlations were significant between Axis 3 and both latitude (F = 21.65, P = 0.002, r = 0.869) and longitude (F = 10.34, P = 0.011, r = 0.775). When species richness (S) was adjusted for sampling effort via rarefaction (Sadj), there was no significant difference for Sadj among spruce–fir sites (χ2 test: P > 0.5). Randomized rarefaction curves do not reach asymptote for any site (Fig. 4). The Figure 4. Randomized rarefaction curves from trap collections for A) spruce–fir sites and B) hardwood sites, based on the Mao Tau richness estimator. Southeastern Naturalist R. Browne, S. Maveety, L. Cooper, and K. Riley 2014 Vol. 13, No. 2 416 slope of the species-accumulation curve for hardwood forest sites was significantly higher than that for spruce–fir sites (slope ± s.e. = 0.027 ± 0.0019 for hardwood sites and 0.0078 ± 0.0056 for spruce–fir sites; F = 3.4, P = 0.021). Abundances of the 12 most common species (n ≥ 45) varied significantly across sites (χ2 tests: P < 0.001 for all 12 species, P < 0.004 with Bonferroni correction). There was no significant difference among Sadj for hardwood sites (χ2 test: P > 0.5). Eight species had sufficient sample size using values adjusted for sampling effort (n ≥ 15, the minimum n for χ2 test for 3 sites with an expected value of n = 5 per site) to test for differences in abundance among sites: Carabus goryi, Cyclotrachelus sigillatus, Dicaelus teter, Pterostichus lachrymosus, Scaphinotus elevatus, Sphaeroderus bicarnitus, and Sphaeroderus stenostomus (P < 0.001 for all species, except P < 0.05 for Dicaelus teter). The only species which did not have a significant difference in abundance among hardwood sites was Sphaeroderus canadensis. The abundance of several genera varied seasonally (Fig. 5). The majority (54.4%) of individuals were trapped in summer, with 23.9% in autumn, 20.2% in spring and 1.5% in winter. Pterostichus was the dominant genus in the summer and fall months, while Sphaeroderus was dominant in winter (composing more than 95% of individuals captured during that season). The other two Cychrine genera, Scaphinotus and Maronetus, showed more restrictive seasonal distributions; we found Scaphinotus only in summer and autumn, and collected the majority (82%) of Maronetus in autumn. At spruce–fir sites, Pterostichus lachrymosus was the mostcommon species for all seasons except winter, when Sphaeroderus stenostomus dominated (Table 4). For hardwood forests, Carabus goryi was the most common species in spring and summer and was relatively abundant in autumn along with Scaphinotus elevatus, and Sphaeroderus bicarinatus was the most abundant in winter. Thus, Cychrines, which prey almost exclusively on gastropods, dominated in winter and autumn at both spruce–fir and hardwood forest sites, although they were Figure 5. Seasonal composition of Carabidae genera from combined spruce–fir sites. Genera consisting of <1.2% of collected individuals are not shown. Southeastern Naturalist 417 R. Browne, S. Maveety, L. Cooper, and K. Riley 2014 Vol. 13, No. 2 collected in extremely low numbers in winter and at just 6 of the 9 spruce–fir sites as well as at all the hardwood sites during that season. Comparison of hand and trap methods for spruce–fir forest sites Of the 39 species collected by hand by Ortiz and Browne (2011), 29 (74%) were also collected in this study. Conversely, 30 (64%) of the 46 species collected in this study were represented in the hand collections. The raw species count collected by trap was larger than that collected by hand, probably due to the higher number of individuals sampled via traps (i.e., sampling effort). However, there was no significant difference (repeated measures ANOVA: P > 0.10) between Sadj, E, and Fisher’s α for our trap collections and the hand collections reported by Ortiz and Browne (2011). While Sadj was not significantly different for trap and hand collections, taxa composition did significantly differ. At the genus level, Gastrellarius was approximately 30 times more likely to be collected by hand than by trap (repeated measures ANOVA: P = 0.003), while Sphaeroderus and Carabus were more likely to be found in traps (repeated measures ANOVA: P = 0.004 and P = 0.047, respectively). In contrast, Pterostichus constituted a large portion of all individuals captured by both trap (49.9%) and by hand (51.2%). The difficulty of separating variation due to technique and due to the different times of collection complicate comparisons between hand and trap techniques, but the data reinforce that while both techniques provide similar diversity estimates, the use of both sampling techniques provide a more complete inventory of the species present. Discussion Although there was no significant difference in Sadj, E, and Fisher’s α between carabid beetles assemblages from spruce–fir sites and hardwood sites, there were marked differences in taxa composition. Pterostichus spp. make up the majority (52%) of individuals sampled at spruce–fir sites, and are significantly less common Table 4. Three most-abundant carabid species by season for combined spruce–fir sites and for combined hardwood sites. Season Spruce–fir Sites Hardwood Sites Summer Pterostichus lachrymosus (31%) Carabus goryi (30%) Carabus goryi (11%) Cyclotrachelus sigallutus (11%) Pterostichus rostratus (10%) Sphaeroderus bicarinatus (11%) Autumn Pterostichus lachrymosus (16%) Scaphinotus elevatus (30%) Sphaeroderus bicarinatus (13%) Carabus goryi (19%) Pterostichus adoxus (11%) Pterostichus lachrymosus (15%) Winter Sphaeroderus stenostomus (78%) Sphaeroderus bicarinatus (53%) Sphaeroderus bicarinatus (11%) Sphaeroderus stenostomus (29%) Sphaeroderus canadensis (9%) Sphaeroderus canadensis (16%) Spring Pterostichus lachrymosus (22%) Carabus goryi (53%) Sphaeroderus canadensis (21%) Sphaeroderus bicarinatus (25%) Sphaeroderus bicarinatus (18%) Sphaeroderus stenostomus (8%) Southeastern Naturalist R. Browne, S. Maveety, L. Cooper, and K. Riley 2014 Vol. 13, No. 2 418 at hardwood sites. Maronetus debilis occurs at a majority of spruce–fir sites but at no hardwood sites, and thus appears to be a strong candidate as an indicator species of spruce–fir forests. We also found Platynus angustatus almost entirely (90 of 92 individuals) in spruce–fir forests and Cylcotrachelus sigillatus almost exclusively (58 of 59 individuals) in hardwood forests. The question of what determines species occurrence at a particular location has been debated for many years. Among the more obvious factors that affect where a species can be found, as suggested in this study, are elevation, temperature, moisture, and forest tree composition (with possible dependence among the factors). Species composition varied by forest type (spruce–fir forests versus hardwood forests) and location (latitude and longitude). Location effects may be related to moisture levels, reinforcing the relationship between geography and rainfall in the southern Appalachians. Large-scale clockwise wind patterns bring moisture into the southern portion of the United States from the Gulf of Mexico, causing precipitation to decrease with latitude in the southern Appalachians (Reinhardt and Smith 2008). Carabid species composition appears to be at least partially influenced by moisture level (Larochelle and Lariviere 2003, Lovei and Sunderland 1996), which in turn might be correlated with elevation. The role of biotic influences such as predator-prey interactions and competition remain largely unknown. We found no significant difference for Sadj among spruce–fir sites. However, species presence varied among sites. Five species were present at all spruce–fir sites; for 12 additional species there were significant differences in abundance among those sites. A question for future studies is whether the species composition is primarily stochastic and follows a neutral model (Hubbell 2001) or has more deterministic components (e.g., niche-assembly theory). Although the number of sample sites for hardwood forests is more limited, the results are similar to spruce–fir sites. Adjusted species richness did not differ among the three hardwood sites, but seven of the eight most common carabid species had heterogeneous abundances among sites. The majority of the most-abundant species were spring breeders that overwinter as adults. These species would be expected to be most active in the spring, summer, and fall months (Larochelle and Lariviere 2003). Cychrines, which can live for more than one year, were present in large numbers in the winter months (Larochelle and Lariviere 2003). In winter, our carabid captures at both spruce–fir and hardwood sites were limited almost entirely to Sphaeroderus. Likewise, at lower-elevation sites in the Piedmont region of North Carolina, Cychrines (primarily Sphaeroderus) dominated overall catch in the winter months (Riley and Browne 2011). Based on the by-catch found in pitfall traps, there is no evidence that Cychrine’s prey (primarily snails) are more common in winter. In fact, more snails have been caught during the summer (R. Browne, unpubl. data). A single year of sampling is not sufficient to reach a definitive conclusion about seasonality in carabid communities. Populations of carabid beetles fluctuate over time, with both the number of individuals collected and the number of species varying by year (Günter and Assmann 2004). Multi-annual collections are needed to definitively determine seasonal patterns. Southeastern Naturalist 419 R. Browne, S. Maveety, L. Cooper, and K. Riley 2014 Vol. 13, No. 2 The integrity of any community comparison is dependent on the degree to which the samples truly reflect the entire community. In this study, species accumulation curves do not reach asymptote, even for sites with high sample numbers. For example, at GSM, where the most individuals were sampled, rarefaction analyses suggest that one new species would be added for approximately 160 new individuals collected. Additional extensive sampling would be needed for a complete inventory of the carabid beetles at all sites. Obtaining a complete inventory often remains an ideal rather than an attainable goal, especially for species-rich taxa in areas with both high alpha and beta diversity such as the southern Appalachians (Carlton and Bayless 2007). Although species accumulation curves are significantly different between hardwood and spruce-fir forest sites, there are little differences in the slope of the curves among hardwood forest sites or in the slope of the curves among spruce-fir forest sites, indicating that species richness is unaffected by altitude, rainfall or tree species composition within a forest type (e.g. among hardwood forest sites). In contrast, the slope of species-accumulation curves increased significantly as elevation decreased in a study in the Peruvian Andes (Maveety et al. 2011). While species richness and number of genera were not significantly different between trap- and hand-sampling techniques, they targeted different taxonomic subsets at both the species and genera levels. Surveys using a combination of both trap and hand collections would more accurately reflect actual species composition (Coddington et al. 1991, Longino et al. 2002, Maveety et al. 2011), and are a necessary pre-requisite for comparing communities across habitats. Acknowledgments We thank the Great Smoky Mountains National Park, Blue Ridge National Park, Pisgah National Forest, Jefferson National Forest, and Mount Mitchell State Park for collection permits. We also are indebted to C. Carlton, V. Bayless, and T. Erwin for assistance in taxonomic identification. Literature Cited Arbogast, B.S., R.A. Browne, P.A. Weigl, and J.G. 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