2013 SOUTHEASTERN NATURALIST 12(2):379–386
Relationships Between Carabid Beetle
Communities and Forest Stand Parameters:
Taxon Congruence or Habitat Association?
Wade B. Worthen1,* and David C.G. Merriman1
Abstract - We sampled ground beetles (Coleoptera: Family Carabidae) at the Furman
Forest in northeast Greenville County, SC, and compared species richness and community
attributes with tree stand richness, diversity, and composition. Beetles were collected
by pitfall trap 1 night/week for 9 weeks from June to August 2011 in 24 plots (200 m2)
varying in tree abundance (11–131 trees/plot), mean tree size (6.5–26.0 cm DBH), species
richness (2–13 species/plot), Simpson’s diversity (1.6–9.1/plot), and composition.
We collected 286 carabids representing 13 genera and 26 species, including a new state
record for South Carolina, Cyclotrachelus hypherpiformis. Carabid abundance, species
richness, and diversity were unrelated to tree abundance, richness, or diversity. However,
carabid abundance and richness were positively correlated with the abundance of
Liriodendron tulipifera (Tulip Poplar), suggesting a preference for mesic habitats. Total
carabid abundance and the abundance and dominance of Carabus goryi (the most abundant
species) were also positively correlated with mean tree size (DBH), suggesting a
preference for older stands. Carabid diversity, abundance, and community structure were
associated with habitat type and stand age (as indicated by dominant canopy species and
tree size) and not canopy tree species richness or diversity.
Species diversity patterns in different taxa are often positively correlated.
Such “cross-taxon congruence” is particularly useful for environmental monitoring
and biodiversity studies, as diversity in one “indicator taxon” may correlate
with environmental pollution, environmental or disturbance gradients, and/or total
biodiversity (Hellawell 1986; Kremen 1992, 1994; Kremen et al. 1993; Noss
1990; Oliver and Beattie 1996). Taxon congruence can occur because of direct
interactions among organisms. Through “bottom-up” relationships, increased
plant or herbivore diversity can spur diversity in consumers and mutualists
(Hunter and Price 1992, Murdoch et al. 1972, Schaffers et al. 2008, Siemann et al.
1998, Southwood et al. 1979). Likewise, through “top-down” forces, increased
predator diversity may reduce competition at lower trophic levels and increase
the diversity of resource taxa (Carpenter et al. 1985, Estes et al. 2011, Letnic et
al. 2012, Paine 1966, Siemann et al. 1998). Congruence can also occur because
diversity in different taxa respond in independent but parallel ways to the spatial
distribution and size of habitat patches (sensu MacArthur and Wilson 1967), to
the same environmental factors (such as temperature, light, water availability, or
a pollutant), or to temporal, “successional” changes in the environment.
1Department of Biology, Furman University, Greenville, SC 29613. *Corresponding author
380 Southeastern Naturalist Vol. 12, No. 2
Ground beetles (Order: Coleoptera, Family: Carabidae) and canopy trees are
likely candidates for congruent groups—they are both diverse and important
components of forest ecosystems. In addition, because most of the >40,000
carabid species are generalist predators (Lövei and Sunderland 1996), their abundance
and diversity might be driven by “bottom-up” forces—by the diversity
and abundance of their invertebrate prey and the diversity of plants their prey
consume. In addition, carabid abundance and diversity increases with leaf-litter
depth (Koivula et al. 1999), soil moisture (Luff et al. 1989), and the presence
of dead wood (Nitterus et al. 2007), which all may correlate with stand age and
canopy tree diversity.
Results testing for congruence between carabids and plants, however, have
been mixed. Carabid diversity correlates with plant diversity across a grassland–
plantation–forest continuum (Fahy and Gormally 1998), but not in grassland
(Vessby et al. 2002) and alpine habitats (Finch and Löffler 2010). In a recent
review, Koivula (2011) concluded that carabids were poor indicators of general
biodiversity. Rather, because carabids may respond more directly to the structural
diversity of plant communities (Brose 2003, Janssen et al. 2009, Sobek et
al. 2009, Taboada et al. 2010), they may be better indicators of habitat type and
environmental conditions. The purpose of this investigation was to test Koivula’s
(2011) conclusions. We compared carabid diversity and community structure
with tree stand diversity and composition across a range of natural forest types in
the southern Appalachians to determine whether carabid communities respond to
canopy tree diversity or habitat type and forest composition.
This experiment was conducted at Furman Forest, a 600-ha tract on the eastern
slope of Hogback Mountain in the northeast corner of Greenville County, SC,
on the first ridge of the Blue Ridge Mountains (383811.84m E, 3894591.25m N).
The forest consists of three adjoining 200-ha parcels owned by The Nature Conservancy
(“Blue Wall Preserve”), The Spartanburg Water System, and the town of
Tryon, NC, and includes two watersheds protected from the ridgeline. There are a
variety of community types on site including the following: ravines with hemlock
(Tsuga canadensis (L.) Carrière [Eastern Hemlock] and T. caroliniana Engelm.
[Carolina Hemlock]); mature stands of Quercus montana Willd. (Chestnut Oak)
and Carya spp. (hickory); mesic hardwood stands of Liriodendron tulipifera L.
(Tulip Poplar) and Acer rubrum L. (Red Maple); dry ridge communities dominated
by Quercus alba L. (White Oak), Q. rubra L. (Northern Red Oak), Q. falcata
Michx. (Southern Red Oak), Q. coccinea Münchh. (Scarlet Oak), Q. velutina
Lam. (Black Oak), and Pinus virginiana Mill. (Virginia Pine); and successional
stands dominated by Virginia Pine and Liquidambar styraciflua L. (Sweetgum).
In 2008, a grid of sampling points was established across the site, with 100 m
between points. Since then, trees have been sampled using circular plots with a
2013 W.B. Worthen and D.C.G. Merriman 381
radius of 8 m (area = 200 m2) centered on each point. The species and size (DBH)
of each tree (>2.0 cm DBH) within each plot were recorded, and species richness
and Simpson’s reciprocal index of diversity measurements (D = 1 / Σ[pi
calculated. For this study, we purposefully selected 24 points that maximized the
range in tree species richness (2 to 13 species per point) and Simpson’s diversity
(1.2 to 9.1). These points also ranged across several habitat types, from successional
pine stands to mesic hardwoods to drier oak/pine ridge communities. A
pitfall trap station was placed at each sampling point, consisting of two 0.95-L
cans buried 1 m apart, and linked by a 0.15- x 1-m aluminum drift fence. Small
holes (≈0.3 cm) were punched in the bottom of each can to prevent rainwater accumulation.
No preservative was used, and the traps were checked after 24 hours
to eliminate the effects of baiting that preservatives and the accumulation of dead
beetles may cause. Although pitfall traps bias samples towards larger, more active
species, they are a common and reliable method for comparing assemblages
of carabids (Spence and Niemelä 1994). For nine weeks between 9 June–5 August
2011, each trap station (n = 24) was left open for one day per week, with
eight different stations opened on each of three consecutive mornings each week.
Cans were covered when not in use. Ground beetles were collected from open
traps each morning after a 24-hr trapping period, euthanized with ethyl acetate,
and identified using Arnett and Thomas (2001), Bousquet (2010), Ciegler (2000),
and Freitag (1969). The abundance of each species, total carabid abundance, species
richness, and Simpson’s diversity were computed for each point (pooling the
data for both cans across all sampling dates). Spearman rank correlation analyses
were used to describe the relationships between carabid communities and tree
communities. Partial correlations were used to describe these relationships while
controlling for variation in tree size.
Results and Discussion
A total of 286 ground beetles were collected from the pitfall traps, representing
13 genera and 26 species (Table 1), including a new state record for South
Carolina, Cyclotrachelus hypherpiformis (Freitag). The communities were
dominated by Carabus goryi Dejean, which accounted for more than 34% of all
individuals. The four species of Carabus (39.3%) and the five species of Dicaelus
(22.1%) accounted for over 60% of the individuals captured (Table 1).
On initial inspection, there appear to be several significant relationships
between canopy tree species richness and carabid communities. Curiously, tree
species richness was negatively correlated with carabid species richness, carabid
abundance, and the abundance of Carabus goryi (Table 2). There were also
relationships with tree size; mean tree DBH was positively correlated with total
carabid abundance and the abundance and dominance (% of total carabids) of
Carabus goryi (Table 2). Because tree abundance and tree species richness were
negatively correlated with mean tree DBH (plots with larger trees containing
fewer individuals and species; Table 2), we hypothesized that the significant
correlations between beetle parameters and tree abundance/richness might be
382 Southeastern Naturalist Vol. 12, No. 2
Table 1. The species of carabid beetles captured at Furman Forest, Greenville County, SC; number
of individuals captured and % representation of each species and genus.
Species n % by species % by genus
Carabus goryi Dejean 97 34.04% 39.30%
Carabus serratus Say 2 0.70%
Carabus sylvosus Say 3 1.05%
Carabus vinctus Weber 10 3.51%
Chlaenius tomentosus (Say) 4 1.40% 1.40%
Cicindela unipunctata Fabricius 32 11.23% 11.23%
Cyclotrachelus hypherpiformis (Freitag) 3 1.05% 12.98%
Cyclotrachelus sigillatus (Say) 34 11.93%
Dicaelus dilatatus dilatatus Say 23 8.07% 22.10%
Dicaelus elongatus Bonelli 1 0.35%
Dicaelus politus Dejean 20 7.02%
Dicaelus purpuratus Bonelli 4 1.40%
Dicaelus teter Bonelli 15 5.26%
Galerita bicolor Drury 2 0.70% 0.70%
Harpalus pensylvanicus (DeGeer) 1 0.35% 1.05%
Harpalus protractus Casey 2 0.70%
Helluomorphoides nigripennis (Dejean) 2 0.70% 0.70%
Pasimachus depressus (Fabricius) 1 0.35% 3.16%
Pasimachus punctulatus Haldeman 8 2.81%
Platynus decentis (Say) 2 0.70% 0.70%
Poecilis lucublandus (Say) 2 0.70% 0.70%
Pterostichus acutipes acutipes Barr 2 0.70% 3.50%
Pterostichus adoxus (Say) 2 0.70%
Pterostichus moestus (Say) 1 0.35%
Pterostichus stygicus (Say) 5 1.75%
Sphaeroderus stenostomus (Weber) 8 2.81% 2.81%
Table 2. Spearman rank correlations between carabid community descriptors and forest stand
parameters at Furman Forest, Greenville County, SC (DBH = diameter at breast height; ns = P >
0.05, * = P < 0.05, ** = P < 0.01, *** = P < 0.001; bold formatting = relationships that remain
significant in partial correlations controlling for DBH (see text for values). One site had no beetles,
so diversity and dominance sample sizes are reduced by 1.
Tulip White Virginia
Tree Tree Tree Tree Poplar Oak Pine
DBH abundance richness diversity abundance abundance abundance
Tree DBH -0.656*** -0.404* -0.150 ns 0.294 ns -0.406* -0.599**
Carabid abundance 0.495* -0.351 ns -0.456* -0.320 ns 0.653*** -0.600** -0.513*
(n = 24)
Carabid richness 0.307 ns -0.339 ns -0.413* -0.248 ns 0.407* -0.480* -0.491*
(n = 24)
Carabid diversity 0.051 ns -0.121 ns -0.188 ns -0.110 ns 0.026 ns -0.048 ns -0.214 ns
(n = 23)
C. goryi abundance 0.711*** -0.658*** -0.465* -0.189 ns 0.514** -0.426* -0.569**
(n = 24)
C. goryi dominance 0.709*** -0.662*** -0.451* -0.158 ns 0.436* -0.287 ns -0.538**
(n = 23)
2013 W.B. Worthen and D.C.G. Merriman 383
spurious. Indeed, when partial correlations were conducted that controlled for
mean tree DBH, there were no statistically significant relationships between
carabid community descriptors and tree abundance, richness, or diversity (P >
0.05). As such, after accounting for variation in tree size, there was no evidence
for “cross-taxon congruence” between carabids and canopy trees.
With respect to habitat type, total carabid abundance, carabid richness, and
C. goryi abundance and dominance were positively correlated with the abundance
of Tulip Poplar and negatively correlated with the abundance of White Oak
and/or Virginia Pine (Table 2). In partial correlations holding mean tree DBH
constant, the positive correlations between Tulip Poplar abundance and carabid
abundance (r = 0.632, df = 21, P < 0.001), C. goryi abundance (r = 0.614, df = 21,
P < 0.01), and C. goryi dominance (r = 0.499, df = 20, P < 0.05) remained significant,
probably indicating a preference for mesic habitats over drier ridges. These
results are consistent with previous research indicating that forest carabids prefer
mesic deciduous stands (Fahy and Gormally 1998; Fuller et al. 2008; Magura
et al. 2000, 2003; Taboada et al. 2010); perhaps because of their preference for
higher soil moisture (Antvogel and Bonn 2001, Luff et al. 1989), more dead wood
(Barton et al. 2009, Fuller et al. 2008, Nitterus et al. 2007) and deeper leaf litter
(Antvogel and Bonn 2001; Fuller 2008; Koivula et al. 1999; Magura et al. 2000,
2005). Because these factors also correlate with stand age and successional stage,
it is not surprising that forest carabid abundance and diversity also correlates
with mean tree size (Tabaoda et al. 2010), as we found in our study. In addition,
the compositional changes we found in C. goryi abundance and dominance with
stand age (as indicated by mean tree DBH) confirmed the results of Jelaska et al.
(2011), who found that the abundance and dominance of large Carabus species
increased through successional time.
Our results also contradict some studies, typically those where a broader range
of habitats were examined. For example, carabid diversity declines with successional
stage and tree size when grasslands or young successional stands are
included (Butterfield 1997, Butterfield et al. 1995, da Silva et al. 2008, Magura et
al. 2001, Niemelä et al. 1996, Silverman et al. 2008). This finding is probably the
result of changes in community composition and a greater diversity of generalists
that prefer open habitats relative to the number of specialized forest species (Butterfield
et al. 1995, da Silva et al. 2008, Fuller 2008, McGeoch 1998, Silverman
et al. 2008).
There are also some discordant patterns regarding habitat preferences of
the dominant species, C. goryi. In our study, C. goryi abundance was strongly
correlated with mean tree DBH and the abundance of Tulip Poplar, suggesting
a preference for older mesic stands. Silverman et al. (2008) also found that
C. goryi was the most abundant species in forest sites in Ohio, but abundance
was greater in ecotonal sites than in forest interior. In Michigan studies, Petrillo
and Witter (2009) found that C. goryi was more abundant in more mesic Sugar
Maple-American Beech sites than in drier Northern Red Oak-American Beech
sites, but Liebherr and Mahar (1979) found that C. goryi was more abundant
in drier White Oak sites than more mesic sites. These discrepancies might be a
384 Southeastern Naturalist Vol. 12, No. 2
function of C. goryi responding to other factors besides moisture and temperature
levels—like leaf-litter depth, dead wood, and stand age—that might vary in different
ways between the sites compared in each study.
In conclusion, carabid beetle abundance and community structure, and the
abundance and dominance of C. goryi, were related to the abundance of canopy
dominants and stand age, probably as a consequence of preferring moist, mesic
habitats with deep leaf litter and lots of dead wood. There was no evidence for
taxon congruence between carabids and canopy trees, once differences in tree
size were taken into account. This was a very small study, however, focusing on
large beetles sampled over a short period. It is certainly possible that a larger,
more extensive survey of all carabids might resolve a pattern of taxon congruence
between these groups.
We thank The Nature Conservancy, Tryon Water District, and Spartanburg Water
Commission for the use of Furman Forest. We thank Janet Ceigler for confirming and
correcting our beetle identifications. We also thank Megan Aprill, Tara Smith, and Amelia
Schulz for their assistance in the field. This research was supported by a grant from the
South Carolina Independent Colleges and Universities.
Antvogel, H., and A. Bonn. 2001. Environmental parameters and microspatial distribution
of insects: A case study of carabids in an alluvial forest. Ecography 24:470–482.
Arnett, R.H., Jr., and M.C. Thomas. 2001. American Beetles. Volume 1. CRC Press, New
York, NY. 443 pp.
Barton, P.S., A.D. Manning, H. Gibb, D.B. Lindenmayer, and S A. Cunningham. 2009.
Conserving ground-dwelling beetles in an endangered woodland community: Multiscale
habitat effects on assemblage diversity. Biological Conservation 142:1701–1709.
Bousquet, Y. 2010. Illustrated Identification Guide to Adults and Larvae of Northeastern
North American Ground Beetles (Coleoptera: Carabidae). Pensoft Publishers, Moscow,
Russia. 562 pp.
Brose, U. 2003. Bottom-up control of carabid beetle communities in early successional
wetlands: Mediated by vegetation structure or plant diversity? Oecologia 135:407–413.
Butterfield, J. 1997. Carabid community succession during the forestry cycle in conifer
plantations. Ecography 20:614–625.
Butterfield, J., M.L. Luff, M. Baines, and M.D. Eyre. 1995. Carabid beetle communities
as indicators of conservation potential in upland forests. Forest Ecology and Management
Carpenter, S.R., J.F. Kitchell, and J.R. Hodgson. 1985. Cascading trophic interactions
and lake productivity. Bioscience 35:634–639.
Ceigler, J. 2000. Ground beetles and wrinkled bark beetles of South Carolina (Coleoptera:
Geadephaga: Carabidae and Rhysodidae). Clemson University, Clemson, SC.
da Silva, P.M., C.A.S. Aguiar, J. Niemelä, J.P. Sousa, and A.R.M. Serrano. 2008. Diversity
patterns of ground-beetles (Coleoptera: Carabidae) along a gradient of land-use
disturbance. Agriculture Ecosystems and Environment 124:270–274.
2013 W.B. Worthen and D.C.G. Merriman 385
Estes,J.A., J. Terborgh, J.S. Brashares, M.E. Power, J. Berger, W.J. Bond, S.R. Carpenter,
T.E. Essington, R.D. Holt, J.B.C. Jackson, R.J. Marquis, L. Oksanen, T. Oksanen,
R.T. Paine, E.K. Pikitch, W.J. Ripple, S.A. Sandin, M. Scheffer, T.W. Schoener, J.B.
Shurin, A.R.E. Sinclair, M.E. Soule, R. Virtanen, and D.A. Wardle. 2011. Trophic
downgrading of planet Earth. Science 333:301–306.
Fahy, O., and M. Gormally. 1998. A comparison of plant and carabid beetle communities
in an Irish oak woodland with a nearby conifer plantation and clearfelled site. Forest
Ecology and Management 110:263–273.
Finch, O.D., and J. Löffler. 2010. Indicators of species richness at the local scale in an
alpine region: A comparative approach between plant and invertebrate taxa. Biodiversity
and Conservation 19:1341–1352.
Freitag, R. 1969. A revision of the species of the genus Evarthus LeConte (Coleoptera:
Carabidae) Quaestiones Entomologicae 5:88–211.
Fuller, R.J., T.H. Oliver, and S.R. Leather. 2008. Forest management effects on carabid
beetle communities in coniferous and broadleaved forests: Implications for conservation.
Insect Conservation and Diversity 1:242–252.
Hellawell, J.M. 1986. Biological Indicators of Freshwater Pollution and Environmental
Management. Elsevier, London, UK. 546 pp.
Hunter, M.D., and P.W. Price. 1992. Playing chutes and ladders: Heterogeneity and the
relative roles of bottom-up and top-down forces in natural communities. Ecology
Janssen, P., D. Fortin, and C. Hébert. 2009. Beetle diversity in a matrix of old-growth
boreal forest: Influence of habitat heterogeneity at multiple scales. Ecography
Jelaska, L.S., V. Dumbović, and M. Kučinić. 2011. Carabid beetle diversity and mean
individual biomass in beech forests of various ages. In D.J. Kotze, T. Assmann, J.
Noordijk, H. Turin, and R. Vermeulen (Eds.). Carabid Beetles as Bioindicators: Biogeographical,
Ecological, and Environmental Studies. ZooKeys 100:287–317.
Koivula, M.J. 2011. Useful model organisms, indicators, or both? Ground beetles (Coleoptera,
Carabidae) reflecting environmental conditions. In D.J. Kotze, T. Assmann,
J. Noordijk, H. Turin, and R. Vermeulen (Eds.). Carabid Beetles as Bioindicators:
Biogeographical, Ecological, and Environmental Studies. ZooKeys 100:287–317.
Koivula, M., P. Punttila, Y. Haila, and J. Niemelä. 1999. Leaf litter and the small-scale
distribution of carabid beetles (Coleoptera, Carabidae) in the boreal forest. Ecography
Kremen, C. 1992. Assessing the indicator properties of species assemblages for natural
area monitoring. Ecological Applications 2:203–217.
Kremen, C. 1994. Biological inventory using target taxa: A case study of the butterflies
of Madagascar. Ecological Applications 4:407–422.
Kremen, C., R.K. Colwell, T.L. Erwin, D.D. Murphy, R.F. Noss, and M.A. Sanjayan.
1993. Terrestrial arthropod assemblages: Their use in conservation planning. Conservation
Letnic, M., E.G. Ritchie, and C.R. Dickman. 2012. Top predators as biodiversity regulators:
The dingo Canis lupus dingo as a case study. Biological Reviews 87:390–413.
Liebherr, J., and J. Mahar. 1979. The carabid fauna of the upland oak forest in Michigan:
Survey and analysis. Coleopterists Bulletin 33:183–197.
Lövei, G.L., and K.D. Sunderland. 1996. Ecology and behavior of ground beetles (Coleoptera:
Carabidae). Annual Review of Entomology 41:231–256.
Luff, M.L., M.D. Eyre, and S.P. Rushton. 1989. Classification and ordination of habitats
of ground beetles (Coleoptera, Carabidae) in northeast England. Journal of Biogeography
386 Southeastern Naturalist Vol. 12, No. 2
MacArthur, R.H., and E.O. Wilson. 1967. The Theory of Island Biogeography. Princeton
University Press, Princeton, NJ. 203 pp.
Magura, T., B. Tóthmérész, and Z. Bordán. 2000. Effects of nature management practice
on carabid (Coleoptera: Carabidae) assemblages in a non-native plantation. Biological
Magura, T., B. Tóthmérész, and T. Molnar. 2001. Forest edge and diversity: Carabids
along forest-grassland transects. Biodiversity and Conservation 10:287–300.
Magura, T., B. Tóthmérész, and Z. Elek. 2003. Diversity and composition of carabids
during a forestry cycle. Biodiversity and Conservation 12:73–85.
Magura, T., B. Tóthmérész, and Z. Elek. 2005. Impacts of leaf-litter addition on carabids
in a conifer plantation. Biodiversity and Conservation 14:475–491.
McGeoch, M.A. 1998. The selection, testing, and application of terrestrial insects as
bioindicators. Biological Reviews 73:181–201.
Murdoch, W.W., C.H. Peterson, and F.C. Evans. 1972. Diversity and pattern in plants and
insects. Ecology 53:819–829.
Niemelä, J., Y. Haila, and P. Punttila. 1996. The importance of small-scale heterogeneity
in boreal forests: Variation in diversity in forest-floor invertebrates across the succession
gradient. Ecography 19:352–368.
Nitterus, K., M. Astrom, and B. Gunnarsson. 2007. Commercial harvest of logging residue
in clear-cuts affects the diversity and community composition of ground beetles
(Coleoptera: Carabidae). Scandinavian Journal of Forest Research 22:231–240.
Noss, R.F. 1990. Indicators for monitoring biodiversity: A hierarchical approach. Conservation
Oliver, I., and A.J. Beattie. 1996. Designing a cost-effective invertebrate survey: A test
of methods for rapid assessment of biodiversity. Ecological Applications 6:594–607.
Ortiz, C.C., and R.A. Browne. 2011. Carabidae (ground beetle) species composition of
southern Appalachian spruce-fir forests. Southeastern Naturalist 10:591–6 08.
Paine, R.T. 1966. Food-web complexity and species diversity. American Naturalist
Petrillo, H.A., and J.A. Witter. 2009. Habitat distribution of carabid beetles (Coleoptera:
Carabidae) in northern hardwood forests of Michigan. Great Lakes Entomologist
Schaffers, A.P., I.P. Raemakers, K.V. Sýkora, and C.J.F. Ter Braak. 2008. Arthropod
assemblages are best predicted by plant species composition. Ecology 89:782–794.
Siemann, E., D. Tilman, J. Haarstad, and M. Ritchie. 1998. Experimental tests of the dependence
of arthropod diversity on plant diversity. American Naturalist 152:738–750.
Silverman, B., D.J. Horn, F.F. Purrington, and K.J.K. Gandhi. 2008. Oil-pipeline corridor
through an intact forest alters ground beetle (Coleoptera: Carabidae) assemblages in
southeastern Ohio. Environmental Entomology 37:725–733.
Sobek, S., I. Steffan-Dewenter, C. Scherber, and T. Tscharntke. 2009. Spatiotemporal
changes of beetle communities across a tree diversity gradient. Diversity and Distributions
Southwood, T.R.E., V.K. Brown, and P.M. Reader. 1979. The relationships of plant and
insect diversities in succession. Biological Journal of the Linnean Society 12:327–348.
Spence, J.R., and J. K. Niemelä. 1994. Sampling ground beetle assemblages with pitfall
traps: The madness and the method. Canadian Entomologist 126:881–894.
Taboada, A., R. Tárrega, L. Calvo, E. Marcos, J.A. Marcos, and J.M. Salgado. 2010.
Plant and carabid beetle species diversity in relation to forest type and structural heterogeneity.
European Journal of Forest Research 129:31–45.
Vessby, K., B. Soderstrom, A. Glimskar, and B. Svensson. 2002. Species-richness correlations
of six different taxa in Swedish seminatural grasslands. Conservation Biology