Biodiversity and Conservation 7, 1589±1605 (1998)

Diversity of litter-dwelling beetles

in the Ouachita Highlands of Arkansas, USA
(Insecta: Coleoptera)
C. E. CARLTON*
Louisiana State Arthropod Museum, Louisiana State University Agricultural Center, Department of Entomology,
Baton Rouge, LA 70803-1710, USA

H. W. ROBISON
Department of Biology, Southern Arkansas University, Magnolia, AR 71753-5000, USA

Received 8 November 1997; revised and accepted 29 March 1998

A survey of forest litter-inhabiting Coleoptera was conducted in deciduous forests of the Ouachita
Mountains in western Arkansas during 1991±1992. A total of 102 Berlese samples were collected,
weighed, and processed during the 12-month study. From 741 kg of sifted forest litter, we counted
and sorted 10 663 adult beetles representing 46 families and 400 species. The family Staphylinidae
was taxonomically and numerically dominant, comprising 46% of species and 63% of individuals.
Problems in assessing species richness of forest litter Coleoptera faunas result from a lack of tax-
onomic revisions, occurrence of sex-limited diagnostic characters, and inadequate information about
larval-adult species associations and life histories. A randomized species accumulation curve indi-
cated that species addition ranged from 14 species per sample during the ®rst ten samples to 1.5
species per sample during the ®nal ten. Richness estimates generated from the empirical data ranged
from 434 species (Michaelis±Menten and Coleman richness estimators) to 590 species (second order
jacknife). The area sampled is biogeographically signi®cant because it harbors numerous habitat-
restricted species that are endemic to the Ouachita Mountains or the Interior Highland region, as
well as disjunct populations of species that are also found in the southern Appalachian Mountains.
Deciduous forest habitats of the Ouachita Mountains, particularly beech-maple riparian forest,
should be given special consideration in forest conservation planning because of their signi®cance as
refugia for these and other endemic and disjunct arthropods.

Keywords: Coleoptera diversity; deciduous forest litter; Ouachita Mountains; endemic and disjunct
species.

Introduction
The recent surge in biodiversity research is beginning to bear fruit in the form of com-
parative surveys of insect faunas, but these surveys are hindered by the incomplete state of
knowledge in insect taxonomy (Samways, 1993), particularly cryptic faunas such as forest
litter and soil arthropods (Hoekstra et al., 1995). Forest litter-inhabiting arthropods are
poorly understood because of their small sizes and cryptic habits. Yet, they are of great
importance to understanding complex processes of nutrient recycling and may contribute
valuable data to studies of comparative biodiversity, forest management, and conservation
(Stork, 1988; Kremen et al., 1993; Hoekstra et al., 1995; Coddington et al., 1997). Species

*To whom correspondence should be addressed.

0960-3115 Ó 1998 Chapman & Hall
1590 Carlton and Robison
composition and regional patterns of diversity of forest litter faunas are still poorly
known, even in areas that have been thoroughly collected, including temperate North
America.
Coleoptera is one of the most diverse groups of organisms represented in forests,
accounting for about 20% of total arthropod diversity (Stork 1988, 1993). Adults of litter-
inhabiting forms can be readily collected and processed for identi®cation and analysis
(Oliver and Beattie, 1993). The state of taxonomy for temperate North American Cole-
optera is reasonably complete, but gaps still exist for some of the more diverse families
represented in forest litter. Despite such problems, Coleoptera are an excellent choice for
comparative biodiversity surveys of forest litter faunas.
The regional focus of this study is the Interior Highlands of western and northern
Arkansas, eastern Oklahoma, and southern Missouri, speci®cally the Ouachita Mountain
region (Thornbury, 1965). Research on the forest litter arthropod fauna of the Interior
Highlands has resulted in the discovery of numerous new species that are endemic to the
Interior Highlands or one of its constituent upland regions. Although taxonomic papers
are an essential ®rst step toward understanding the composition and faunal anities of
forest litter Coleoptera of the Interior Highlands, they have not provided a comprehensive
picture of the total diversity from any single habitat in the region. Such studies are
essential in identifying regional and continental-scale patterns of diversity across large
taxa that can be applied to questions of biodiversity, conservation management, and other
avenues of inquiry that require landscape-scale information about the biota.
In this paper, we report the results of a study of the litter-dwelling Coleoptera fauna of
deciduous forest habitats in the Ouachita Mountains in west-central Arkansas (Fig. 1).
The two habitats sampled are characterized by di€erent stand characteristics, but grade
into each other and represent the range of predominantly deciduous habitat represented in
the Ouachita Mountains. We chose this area for detailed study because beech-maple
forests in this area are similar in ¯oral composition to cove or mixed mesophytic forests of
the southern Appalachians (Braun, 1950). Collections from these habitats have yielded
localized endemic and disjunct species of Coleoptera that reinforce this biogeographic
association. We provide a species-level analysis of Coleoptera from 102 Berlese samples
collected during a 12-month period during 1991±1992, including a list of most abundant
taxa, rank abundance analysis, species accumulation curves, richness estimates, and in-
formation about seasonal abundance and habitat speci®city. Finally, we focus on selected
endemic and disjunct species and discuss their ranges and habitat associations.

Methods
North-facing mesic oak-hickory forest and moist beech-maple riparian forest were sam-
pled in the Ouachita Mountains of western Arkansas (Fig. 1). Six localities were sampled
six to 12 times during the study, usually at monthly or semi-monthly intervals (solid
symbols in Fig. 1). Ten additional localities were sampled once (open symbols in Fig. 1).
The following habitat descriptions are based in information provided by Bates (1989).

Mesic oak-hickory forest. In the Ouachita Mountains these habitats occupy rocky, north-
facing slopes with little or no direct sunlight penetration during the growing season. Floral
communities are dominated by white oak (Quercus alba), black oak (Q. velutina), northern
red oak (Q. rubra), and mockernut hickory (Carya tomentosa). Understory trees include
Litter beetle diversity in Arkansas 1591

Figure 1. Outline map of Arkansas showing major subdivisions of the Interior Highlands, location
of study area (inset), and sample localities. Triangles indicate oak-hickory sites, circles indicate
beech-maple sites. Solid symbols indicate regularly sampled sites, open symbols indicate sites sam-
pled once.

redbud (Cercus canadensis), ¯owering dogwood (Cornus ¯orida), hop hornbeam (Ostrya
virginiana), white ash (Fraxinus americana), black cherry (Prunus serotina), and American
holly (Ilex opaca). Growth of woody vines is often vigorous, represented mainly by poison
ivy (Toxicodendron radicans) and Virginia creeper (Parthenocissus quinquefolia).

Beech-maple riparian forest. This habitat is common along watercourses that are not
regularly inundated by high water. Dominant tree species are American beech (Fagus
grandifolia) and red maple (Acer rubrum). Other common tree species associated with this
community are sweetgum (Liquidamber styraci¯ua), sycamore (Platanus occidentalis),
1592 Carlton and Robison
sugarberry (Celtis laevigata), and American elm (Ulmus americana). Many of the same
understory species found in mesic oak-hickory forest also occur in beech-maple riparian
forest, but umbrella magnolia (Magnolia tripetala) is a common and distinctive compo-
nent. In addition to poison ivy and Virginia creeper, common vines include greenbriers
(Smilax spp.), wild yam (Dioscorea sp.), rattan vines (Berchemia scandens), and grapevines
(Vitis spp.). This community type is diverse and re¯ective of the rugged nature of this
portion of the Ouachita Mountains. Deep ravines and areas of pronounced relief are
common. Each drainage supports di€erent combinations of plant species, depending on
light availability, soil texture and composition, local geology, and other physical and biotic
variables. The similarity of these riparian habitats to the mixed mesophytic forests of the
southeastern United States, particularly the southern Appalachians, was discussed by
Braun (1950).
These descriptions identify di€erences in stand characteristics between the two forest
types in their purest forms. However, the two are contiguous within the study area and the
boundaries are rarely clearcut. Pure examples of beech-maple forest are found in narrow
zones adjacent to continuously-¯owing streams and hillside seeps, but upland species
begin to appear immediately upslope. The more moisture dependent beech-maple com-
ponents disappear quickly on ¯at hillsides and on drier, southern exposures, but may
persist further upslope in small ravines, seeps, and other areas of persistent moisture. Also,
a number of tree species are found in both types of habitat, di€ering only in relative
abundance.

Sampling and processing protocol. Litter samples were collected by sifting moist organic
litter on the forest ¯oor, including thoroughly rotted logs, leaf packs, root mats, and ¯ood
debris, through a 1 cm mesh wire sifter. The sifted sample was weighed and held in a cloth
bag at room temperature until extraction. Organisms were extracted using Berlese funnels
equipped with 100 W incandescent lights for at least 48 h. Berlese residues were preserved
in 70% ethanol prior to processing.
Coleoptera were sorted from each sample, and examples of species from each sample
were mounted and labelled. If multiple specimens of a species were present in a given
sample, a tag was axed to the specimen noting the number of specimens of that species
that were not mounted. If there was doubt about whether a series of specimens belonged
to the same species, all were mounted and labelled. Species were identi®ed or sorted to
presumed same-species series and assigned morphospecies numbers, as elaborated under
Results. All morphospecies sorting was done by the ®rst author. In addition to determi-
nations by the ®rst author, numerous specialists provided identi®cations for taxa within
their areas of expertise (see Acknowledgements). Specimens are deposited in the Univer-
sity of Arkansas Arthropod Museum (Fayetteville, AR) and the Louisiana State
Arthropod Museum (Baton Rouge, LA).

Data analysis. Species accumulation curves and richness estimates were generated using
EstiMateS Richness Estimator Program, Version 4.1 (R. K. Colwell, unpublished). Col-
well's Richness Estimator Program, EstiMateS, generated estimates of species richness
based on empirical data using published formulae that have been adapted from theory to
this purpose (R. K. Colwell, unpublished). The program generates nine richness estimator
curves from each data set. The Coleman richness expectation was based on species±area
relations across di€erent patch sizes (Coleman et al., 1982). The Michaelis±Menten equa-
Litter beetle diversity in Arkansas 1593
tion is a two-parameter hyperbolic function used in enzyme kinetics. Raaijmakers (1987)
used maximum likelihood to estimate parameters and their variances for the Michaelis±
Menten equation and this approach was adopted by Colwell and Coddington (1994) to
estimate species richness. Chao 1 (abundance-based), Chao 2 (incidence-based), and inci-
dence-based coverage (ICE) richness estimators were developed to estimate population
sizes from capture±recapture data (Chao, 1984, 1987; Lee and Chao, 1994). The abun-
dance-based coverage estimator (ACE) was developed to establish stopping rules in com-
puter debugging by analysing the frequency of failures occurring at unequal rates (Chao
et al., 1993). First and second order jackknifes and bootstrap richness estimators are non-
parametric incidence-based methods for estimating the number of species under quadrat
sampling (Smith and van Belle, 1984). The methodology and derivation of the estimators,
and the e€ect of di€erent variables on their performances were described in detail by
Chazdon et al. (1997), Cowell and Coddington (1994) and Coddington et al. (1997).
Samples were added randomly until all had been included in the calculations and this
process was repeated for the prescribed number of randomizations. One hundred ran-
domizations and ten abundance classes were included in the analysis. In cases where
species determinations were possible only for males, mixed series of undetermined females
were excluded from the richness estimator analysis. The result was a series of smooth
richness estimate curves that could be compared with each other and with the species
accumulation curve.
Abundances for pooled samples are expressed as total numbers of individuals collected
per species, numbers of specimens of each species per kg of sample and abundance
ranking. Numbers of individuals and individuals per kg of sample were also broken down
by habitat for species showing a high degree of habitat speci®city and represented by ten
or more individuals in the pooled samples. This ®gure represented our best estimate of an
adequate sample size.

Results
Species richness and abundance. We processed 741 kg of sifted forest litter during 12
months of collecting, 295 kg of litter from oak-hickory forest and 446 from beech-maple.
Samples averaged 6.75 kg (range, 2.27±16.86: SD 1.560). We sorted 10 663 beetles into 46
families and 400 species. We named 185 species and sorted the remaining 215 to mor-
phospecies, generally within named genera. A complete species list, broken down by
specimens kg)1, is available from the ®rst author. The family Staphylinidae accounted
for 43% of total species and 63% of individuals (170 and 6686, respectively). The
Scydmaenidae was a distant second, accounting for 8% of species and 8% of individuals
(30 and 808), followed by Curculionidae, with 6% of species and 4% of individuals
(25 and 386) and Carabidae, with 5% of species but less than 1% of individuals (21 and
91). The superfamily Staphylinoidea contributed 57% of the total species richness (227
species).
Conoplectus canaliculatus (Brendel) (Staphylinidae: Pselaphinae) was the most abundant
species in the survey, represented by 990 individuals, or slightly less than 10% of the total
beetles counted. The ten most abundant species accounted for approximately one-third of
the total individuals (3558 of 10 664) (Table 1). At the other end of the abundance scale,
191 species (47% of total) were represented by three or fewer individuals, and 119 species
(29% of total) were represented by a single individual (Fig. 2).
1594 Carlton and Robison
Table 1. Ten most abundant species in the pooled data set

Abundance rank Species n Individuals per kg of litter

1 Conoplectus canaliculatus 990 1.33

2 Pteryx sp. 570 0.77
3 Leptura sp. 3 510 0.69
4 Philonthus alumnus 446 0.60
5 Carpelimus sp. 438 0.59
6 Trimiomelba dubia 266 0.36
7 Ptinella sp. 223 0.30
8 Sepedophilus sp. 3 212 0.29
9 Leptoplectus pertenuis 207 0.28
10 Agathidium sp. 2 206 0.28

Species accumulation through 100 randomizations of the sample sequence gave a

smooth curve with little change in slope beyond about the 30th sample (Fig. 3). New
species were accumulated at mean rates of 14 species per sample during the ®rst ten
samples, 2.4 species per sample between 45 and 55 samples, and 1.5 species per sample
during the ®nal 10 samples. The second-order jackknife richness estimator produced the
highest prediction of 590 species (Fig. 4). Both the Coleman richness expectation curve
and the Michaelis±Menten richness estimator (based on means) gave the most conserva-
tive estimates of 434 species (Michaelis±Menten shown in Fig. 4). The remaining esti-
mators fell between these values, as follows: bootstrap, 454; Chao 1, 496; abundance-base
coverage estimator, 506; ®rst order jacknife, 525; Chao 2, 528; and incidence-based cov-
erage estimator, 534.

Figure 2. Rank abundance distribution (log scale) for Ouachita forest litter Coleoptera.
Litter beetle diversity in Arkansas 1595

Figure 3. Species accumulation curve through 100 randomizations.

Seasonal abundance. Bias caused by uneven monthly sampling e€ort prevented reliable
direct measurement of seasonal patterns of species richness. However, seasonal variation
in Coleoptera abundance corrected for sample size provided an indirect method of esti-
mating seasonal variations in species richness. This approach assumes that species richness
is positively correlated with individual abundance. Abundance was seasonally bimodal
when measured as numbers of individuals per kg of litter, with peak abundances in March
(19 individuals kg)1) and October (21 individuals kg)1) (Fig. 5). Minimum abundances
occurred in June (11 individual kg)1) and January (10 individuals kg)1).
Habitat associations. Thirty-three species showed an association to one or the other
habitat in our data set (Table 2). Criteria used to select these species included 75% or
more of individuals (measured by individuals per kg of litter) in one habitat and repre-
sented by ten or more specimens in the pooled data (i.e a reasonable sample). Seventeen
species showed bias in favour of beech-maple litter habitat and 16 in favour of oak-
hickory.

Discussion
Species richness and abundance. Two sources of error probably introduce a conservative
bias in our empirical species richness and abundance counts. First, as pointed out by
Spence and NiemelaÈ (1994), our sampling technique might have resulted in underrepre-
sentation of large, active species, such as many carabids, that were able to take evasive
action during our approach. Second, imprecision in sorting morphospecies is likely to
result in lumping of similar appearing species rather than splitting single species into two
1596 Carlton and Robison

Figure 4. Curves generated by EstiMateS (R. K. Colwell, unpublished) showing maximum

(Jack2 ˆ second order jacknife) and minimum (MMMeans ˆ Michaelis±Menten by means) esti-
mates of species richness based on the empirical data.

or more. Oliver and Beattie (1996) noted that when sorting was performed by specialists
these lumping events had minimal e€ect on the overall outcome.
Some taxa could not be as thoroughly sorted or precisely identi®ed as others. For
example, featherwing beetles (Ptiliidae) were abundant in the samples, and 1159 were
counted and included in the database. However, spot checks of residues from which
presumably all beetles had been removed revealed that a few remained in each sample.
Featherwing beetles are among the smallest beetles, ranging in length from 0.50 to
1.00 mm, and are easily overlooked. Further, the presence of numerous polymorphic
species that varied in the development of wings and eyes, combined with a lack of usable
keys, made sorting and identi®cation of species problematic at best. Ptiliids are therefore
underrepresented in our database numerically and, most likely, taxonomically.
Taxonomic diculties were most pervasive at the generic level within the staphylinid
subfamily Aleocharinae. A combination of large numbers of undescribed taxa and no-
menclatural instability render this subfamily one of the most dicult groups to deal with
in biodiversity surveys of Coleoptera. Additional problems of naming species of Ale-
ocharinae occur in many genera because revisionary studies and/or regional identi®cation
guides are lacking. Within the family Scydmaenidae, the genera Connophron and Euconnus
were represented in our data set by 24 recognizable species, but only three species could be
named by specialists. Within Connophron, eight of the ten species were con®rmed as
undescribed. A revision of the genus is underway (K. Stephan, pers. comm.). On a positive
Litter beetle diversity in Arkansas 1597

Figure 5. Seasonal abundance expressed as number of individuals of all species per kg of sifted litter.

Table 2. Species showing 75% or greater bias to one litter habitat (by individuals per kg of litter and
percentages in each habitat) and represented in pooled samples by ten or more specimens

Ind. per kg (%) in Ind. per kg (%) in

Taxa n oak-hickory litter beech-maple litter

Carabidae
Anillinus sp. 10 0.007 (18) 0.031 (82)
Ciidae
Cis punctulatus 17 0.047 (87) 0.007 (13)
C. miles 11 0 (0) 0.025 (100)
Corylophidae
Bathona sp. 1 13 0.044 (100) 0 (0)
Gronevus sp. 1 17 0 (0) 0.038 (100)
Cryptophagidae
Cryptophagus dicilis? 19 0.064 (100) 0 (0)
Curculionidae
Lymantes sandersoni 55 0.020 (17) 0.120 (83)
Derodontidae
Derodontus esotericus 11 0.034 (94) 0.002 (6)
Endomychidae
Lycoperdina ferruginea 12 0.041 (100) 0 (0)
Clemmus minor 11 0 (0) 0.247 (100)
Histeridae
Onthophilus de¯ectus 15 0.051 (100) 0 (0)
1598 Carlton and Robison
Table 2. (Continued)

Ind. per kg (%) in Ind. per kg (%) in

Taxa n oak-hickory litter beech-maple litter

Ptiliidae
Ptiliolum sp. 1 25 0.074 (91) 0.007 (9)
Staphylinidae
Aleocharinae genus sp. 1 21 0.003 (6) 0.045 (94)
Aleocharinae genus sp. 5 32 0.105 (98) 0.002 (2)
Aleocharinae genus sp. 13 18 0.054 (92) 0.004 (8)
Euaesthetinae
Stictocranius puncticeps 24 0.071 (91) 0.007 (9)
Osoriinae
Osorius planifrons 12 0 (0) 0.027 (100)
Paederinae
Achenomorphus corticinus 79 0.030 (16) 0.157 (84)
Homaeotarsus sp. 15 0.007 (19) 0.029 (81)
Lathrobium sp. 21 0.071 (100) 0 (0)
Sciocharis carolinensis 163 0 (0) 0.365 (100)
Proteinae
Megarthrus sp. 38 0.115 (93) 0.009 (7)
Pselaphinae
Rhexius schmitti 17 0.056 (100) 0 (0)
Ouachitychus parvoculus 21 0.003 (6) 0.045 (94)
Scaphidiiniae
Baeocera apicalis 11 0.030 (88) 0.004 (12)
B. charybda 15 0.041 (85) 0.007 (15)
B. scylla 10 0.034 (100) 0 (0)
Scaphisoma pusillum 19 0 (0) 0.043 (100)
S. terminatum 13 0 (0) 0.029 (100)
Staphylininae
Diochus schaumi 78 0.244 (95) 0.013 (5)
Nudobius luridipennis 20 0.003 (15) 0.043 (85)
Tachyporinae
Bryoporus sp. 2 12 0.007 (24) 0.022 (76)
Sepedophilus sp. 2 13 0 (0) 0.023 (100)

note, the recent development of internet websites dealing with some taxa has made useful
taxonomic information more accessible (e.g. the University of Kansas Snow Entomologi-
cal Museum's aleocharine website at http://www.nhm.ukans.edu/ksem/peet/peet1.htm/,
J. S. Ashe, pers. comm.).
A troubling source of error in gathering precise quantitative data at the species level
was caused by species determinations that relied on the availability of the male sex. This
problem was particularly relevant to the identi®cation of staphylinids and scydmaenids.
Multiple species within a single genus were often present in a sample, but only males could
be separated. Approximately 50 species (13% of total species) fell into this category.
Within these 50 species, there were approximately 500 female individuals (5% of total
individuals) that could not be sorted to species. This may have a€ected the richness
estimators by arti®cially in¯ating the number of species present as uniques or small
Litter beetle diversity in Arkansas 1599
numbers of individuals, since females could not be included in these ®gures. We know of
no way to adequately account for this discrepancy at present, and it will continue to be a
problem in analysing diversity patterns in faunas with species that can only be identi®ed
using sex-limited characters.
Do surveys such as this provide a complete picture of species richness in a habitat such
as forest litter? The predictive power of the estimators is only beginning to be assessed, and
long-term resampling e€orts are needed to determine which of these provide the most
accurate predictions under a given set of circumstances. The performances of most of these
estimators were analysed by Chazdon et al. (1997) using species±area data from woody
plants in Costa Rica. In their study the Jackknife estimators produced high estimates, as
was the case in our study. Michaelis±Menten gave low estimates of species richness in their
study (though Chao 1 gave the lowest), which is consistent with our results, in which it
gave the lowest estimate. Chazdon et al. (1997) favoured incidence-based estimators in
samples containing a large number of patchily distributed species, with ICE as the fa-
voured method for highly diverse data sets. Because our data seem to ®t this description,
the ICE ®gure of 534 may be the best estimate of species richness in the present study.
The addition of species at low frequency would likely continue through a great number
of samples during extended sampling of forest litter, which constitutes a subset of the total
habitat complex of the forest. Apparently rare, patchily distributed species that are per-
manent residents of forest litter would be added at a low rate and would be of direct
interest to studies of that habitat. In theory, such species would eventually all be sampled,
and the species accumulation curve would approach an asymptote. However, it is likely
that accidental species (tourists) from elsewhere in the forest and species that only spend a
portion of their life cycles in the litter would be added for a much longer sampling period,
thus obscuring the point at which permanent residents had been thoroughly sampled.
Examples of temporary residents include many adult Chrysomelidae and Curculionidae,
which utilize litter as shelter during harsh weather, and ptylodactylids, which utilize it
during larval development. The detritivorous larvae of Ptylodactyla spp. were abundant in
litter samples during this survey, but the adults, which are mainly arboreal, were rarely
collected. To gain a complete picture of Coleoptera species richness and relative abun-
dance in forest litter, larval forms need to be included in analyses. At present, this
probably is feasible only at family and subfamily levels for most taxa. Obviously, more
information needs to be gathered about biologies of the component taxa at various life
stages in order to gain a more complete picture of how these assemblages interact.

Seasonal abundance. The spring-fall peaks in Coleoptera abundance (Fig. 5) were con-
sistent with anecdotal observations of patterns of terrestrial insect activity and diversity in
Arkansas by us. However, the March peak was earlier than usually observed in non-litter
habitats, where peak activity occurs during late May and early June. Mid-summer patterns
were strongly in¯uenced by ¯uctuations in precipitation in the region. The January min-
imum may be an underestimate if many diapausing adults were not recovered during
Berlese extraction. Studies in which equal volumes of litter are processed during each
sampling interval would allow a more direct measurement of seasonal diversity patterns
among forest litter arthropods.

Habitat associations. The two deciduous habitats di€er in co-dominant tree species and
understory vegetation, presumably due to abiotic di€erences such as soil depth, soil
1600 Carlton and Robison
moisture content and retention, and perhaps local microclimate. Despite the stand and
topographic di€erences, the vast majority of commonly collected species were found in
both habitats, suggesting that partitioning in the litter fauna was not correlated to that of
the dominant plant cover. Nineteen of the 33 species that showed habitat speci®city were
fungivores. Fungi are ephemeral and patchily distributed resources. Thus, at least some of
the apparent habitat speci®city suggested in Table 2 is probably due to patchily distributed
hosts that may or may not be habitat related. Again, only by knowing individual species
biologies can conclusions be drawn about whether patterns are correlative or causal.

Endemic and disjunct species. The Interior Highlands (Fig. 1) consist of several moun-
tainous regions approximately 375 000 km2 located in Arkansas, eastern Oklahoma, and
southern Missouri. The Ozark Mountains are a series of severely eroded fault block
plateaus, and the Ouachita Mountains are a series of east±west trending anticlinal ridges.
The two mountain ranges are separated by an elongate structural feature called the
Arkoma Basin through which the Arkansas River ¯ows. The Ozarks formed as a result of
a localized bulge in the Earth's crust. The Ouachitas formed as a result of tectonic in-
teractions between crustal plates (Wickham et al., 1976). Virtually all mountain-building
activity had ceased by the end of the Carboniferous, some 300 million years ago
(Thornbury, 1965). The independent origins and subsequent isolation of the Ozarks and
Ouachitas by the Arkoma Basin is an important but poorly understood factor a€ecting the
distributions of relict species within the Interior Highlands.
We are aware of at least nine species of forest litter Coleoptera that are endemic to the
Ouachita Physiographic Region or represent disjunct populations of taxa that are also
found in the southern Appalachians (Table 3). At least two species of blind, wingless
Anillinus ground beetles are found in the Ouachita Mountains, with a single occurrence in
the Ozark Mountains (Fig. 6). It was not possible to identify species because no reliable
key to species or revision was available. Thus, we cannot say with certainty whether these
species are endemic to the region or represent species having disjunct distributions. One of
the two species was strongly associated with beech-maple litter (14 of 16 individuals).

Table 3. Coleoptera apparently endemic to or having disjunct populations in the Ouachita

Highlands of Arkansas and Oklahoma

Endemic (E)
Taxa or disjunct (D) References

Carabidae
Anillinus spp. (two species)
Scaphinotus parisiana
Staphylinidae
Arianops copelandi
A. sandersoni
A. stephani
Brathinus nitidus
Ouachitychus parvoculus
Pseudactium magazinensis
Tychobythinus carolinus
? This paper (Fig. 6)
E Allen and Carlton (1988)
E Carlton and Cox (1990)
E Carlton and Allen (1989)
E Carlton and Allen (1989)
D This paper (Fig. 6)
E This paper (Fig. 7)
E Carlton and Chandler (1994);
this paper (Fig. 7)
D This paper (Fig. 7)
Litter beetle diversity in Arkansas 1601

Figure 6. Arkansas localities for Anillinus spp. (circles) and Brathinus nitidus (triangles).

Twelve described and a number of undescribed species of Anillinus are known from the
eastern United States. All but two of the described species occur in the southern Appa-
lachian Mountains. One species occurs in central Texas, another in Indiana (Erwin et al.,
1997).
Previously unrecorded disjunct populations of the staphylinids Brathinus nitidus
LeConte (Omaliinae) and Tychobythinus carolinus (Casey) (Pselaphinae) were discovered
during the study. One specimen of B. nitidus collected by us and an additional specimen
collected by R. Leschen from Mt Magazine (Logan County) represent a substantial range
extension (Fig. 6). The species is documented from southeastern Canada, northeastern
United States, and the Appalachian Mountains as far south as northern Alabama (Peck,
1975). It is apparently an obligate cavernicole in the southern portions of its range to the
east, but Arkansas specimens were collected from upland sphagnum bogs and moist
riparian litter in beech/maple forest. The biology and larval stages of B. nitidus were
described by Thayer (1985). The only other known populations of T. carolinus occur in the
vicinity of Asheville, NC, which is the type locality. Though our samples contained only
two specimens, the species can be readily collected in litter around the margins of
sphagnum bogs in the Ouachita Mountains, which are scattered throughout moist valleys
(Fig. 7).
1602 Carlton and Robison

Figure 7. Arkansas localities for Ouachitychus parvoculus (circles), Pseudactium magazinensis

(squares) and Tychobythinus carolinus (triangles).

The pselaphine staphylinids Ouchitychus parvoculus Chandler and Pseudactium maga-

zinensis Carlton and Chandler are highly localized endemics in the Ouachita Highlands.
O. paravoculus was previously known only from the male holotype collected on Mt
Magazine (Chandler, 1988). We collected 21 specimens from three localities (Fig. 7): 20
were found in beech-maple litter and one in oak-hickory, Chandler (1988), in his cladistic
analysis of genera of the tribe Tychini, concluded that Ouachitychus and Cylindrarctus are
sister taxa. The latter genus is widely distributed in eastern North America and is sym-
patric with Ouachitychus in the Ouachitas. P. magazinensis was described by Carlton and
Chandler (1994) based on a single specimen collected on Mt Magazine and two collected
during this survey (Fig. 7). A second Arkansas species occurs in the Ozark Mountains to
the north (Carlton, 1995). Eight of the nine remaining species are found in the southern
Appalachians and in relict mixed mesophytic habitats along the Gulf Coastal Plain. One
species was described based only on the holotype collected in Iowa.
The remaining species listed in Table 3 were not represented in the collections on which
this study is based, but represent additional endemic taxa and reinforce the uniqueness of
the Ouachitas as an area of endemism. One of our expectations entering the study was that
additional species of the pselaphine genus Arianops would be discovered, but this did not
happen. The three species located in the Ouachitas form a well-supported monophyletic
Litter beetle diversity in Arkansas 1603
clade (Carlton, unpublished). Thirty additional described species are restricted to the
southern Appalachians and a single undescribed species occurs on the Gulf Coastal Plain
in Alabama.
Additional species that are endemic to the Interior Highlands but not restricted to the
Ouachita Mountains were represented in our collections. Metophthalmus sandersoni
Andrews (Lathridiidae), Conoplectus susae Carlton (Staphylinidae: Pselaphinae), and
Oopterinus distinctus O'Brien (Curculionidae) are all restricted to the uplands of northern
and western Arkansas, eastern Oklahoma and southern Missouri. Derops divalis (San-
derson) (Staphylinidae: Tachyporinae) is largely restricted to this area but also extends
into the easternmost fringes of the Ozarks in southern Illinois.
The list of endemic and disjunct organisms in the Ouachita and Ozark Highlands is
long, and re¯ects the long history of the region as an emergent landform and refugium. In
the Ouachita Highlands, the beetles discussed above are joined by many other endemic
invertebrates and a few vertebrates. Examples include an earthworm, Diplocardia meansi
Gates, endemic to Rich Mountain (Polk County), a terrestrial snail, Mesodon magazinensis
(Pilsbry and Ferriss), endemic to Mt Magazine (Logan County) and listed as threatened
under the Endangered Species Act, and numerous species of amphipods, myriopods and
insects. Vertebrates include a cat®sh, the Caddo madtom, Noturus taylori Douglas, the
palebacked darter, Etheostoma pallididorsum Distler and Metcalf, ®ve species of Plethodon
salamanders and one species of Desmognathus. For a detailed treatment of endemic
organisms in Arkansas, see Robison and Allen (1995).
In his discussion of the evolution of Plethodon lineages Highton (1995) cited Pliocene
climatic drying and reduction of low elevation forests as a mechanism for isolation and
subsequent speciation of restricted montane species, including the Ouachita endemics and
many of their southern Appalachian relatives. This explanation probably accounts for
some of the endemic species of beetles and other arthropods that show phylogeographic
congruence with the salamanders. But, a multilayered chronology of events has probably
a€ected the isolation of Ouachita endemics and disjuncts. Possible older events include
pre-Tertiary upland connections with the Appalachians as discussed by Carlton and Cox
(1990) for Arianops spp. Younger events include the often invoked climate cycles of the
Pleistocene, cited by Ross (1965) as causing shifts in the distributions of caddis¯ies and
stone¯ies that resulted in the presence of endemic and disjunct species in the Ouachitas
and Ozarks.

Conclusions
This study serves as a starting point for comparing species richness and abundance of
forest litter habitats in the Ouachita Mountains with those in other regions and for
assessing future changes in forest litter faunas within the Ouachita Mountains due to
anthropogenic or natural changes. The picture is one of a rich but still relatively poorly
understood and taxonomically challenging Coleoptera community, comprising between
400 and 600 species. We o€er a baseline analysis that can be used to examine the per-
formance of several species richness estimators currently being employed to predict spe-
cies diversity on the basis of a limited number of samples. Interpretation of studies such as
this is compromised by diculties in identi®cation of species due to our incomplete
knowledge of species diversity within speciose but little-known taxa, the reliance of
identi®cation on sex-limited characters, our inability to identify larvae at lower taxonomic
1604 Carlton and Robison
levels, and the lack of natural history information about the majority of species. These
problems are serious barriers even in relatively well studied regions such as the eastern
United States.
In terms of future research and regional conservation planning, we point to the oc-
currence of a number of endemic and disjunct species associated with the beech-maple
habitats we sampled and suggest that these areas receive special attention as refugia for
relict populations. The occurrence of endemic species and disjunct populations within the
Coleoptera and other organisms is an indication that many additional endemic species
await discovery within other taxa of Ouachita forest litter invertebrates.

Acknowledgements
We are grateful to the following coleopterists for providing or con®rming identi®cations:
Robert S. Anderson (Canadian Museum of Nature), James S. Ashe (Snow Entomological
Museum, University of Kansas), Sean O'Keefe (Univeristy of California, Berkeley),
Richard Leschen (Landcare Research New Zealand), Alfred Newton Jr (Field Museum of
National History), Karl Stephan (Red Oak, OK), and Margaret Thayer (Field Museum of
Natural History). We thank R.K. Colwell (University of Connecticut, Storrs) for making
the EstiMateS Species Richness Estimator Program available. This project was supported
in part by a grant from the Louisiana Education Quality Support Fund to C. Carlton and
D. Prowell. Field work was supported by a Challenge Cost Share Agreement with the US
Forest Service, Ouachita National Forest. We thank Janice Bossart, Julie Denslow,
Deborah Landau, Richard Leschen, Dorothy Prowell, John Spence, and one anonymous
reviewer for their reviews of the manuscript.

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