1Louisiana State Arthropod Museum, Department of Entomology, Louisiana State
University Agricultural Center, Baton Rouge, LA 70803. *Corresponding author -
Documenting Beetle (Arthropoda: Insecta: Coleoptera)
Diversity in Great Smoky Mountains National Park:
Beyond the Halfway Point
Chris Carlton1,* and Victoria Bayless1
Abstract - The current Coleoptera (beetle) Taxonomic Working Group of the All
Taxa Biodiversity Inventory (ATBI) has been active since June 2001. It consists of a
core group of students and researchers, headquartered at the Louisiana State Arthropod
Museum (LSAM), Baton Rouge, LA and is supported by a network of 42 specialists
worldwide. Our starting point was ≈700 species based mainly on specimens
collected prior to 2001 and deposited in the Great Smoky Mountains National Park
(GSMNP) museum. Our current beetle count for GSMNP is 2131 species from 103
families. Using periodic species accumulation analyses plus independent work by
cooperators, we have documented ≈1400 new species records for the park and 16,370
specimen records in our Biota® relational database. This includes 42 species new to
science. We predict the total beetle diversity for the park will be ≈3000 species based
on extrapolation of data for the 15 largest families in eastern North America. Most
additional records are expected from taxonomic work on the following families:
Staphylinidae (rove beetles), Curculionidae (weevils, bark, and ambrosia beetles),
Tenebrionidae (darkling beetles), Scarabaeidae (scarab beetles), Latridiidae (minute
scavenger beetles), and Ptiliidae (feather-winged beetles). Progress has been
hampered by a lack of taxonomic expertise and logistical problems associated with
large specimen volumes. The former problem is society-wide and the latter has been
partially solved. The vast majority of specimens processed to date were derived from
structured protocols conducted during the initial phase of the ATBI. Current and future
efforts will be focused on samples collected during organized beetle bioblitzes
and/or using specialized techniques targeting taxa that cannot be obtained using
The most recent treatments of American beetles (Arnett and Thomas 2001,
Arnett et al. 2002) lists 131 families in the US, of which 103 are known from
Great Smoky Mountains National Park (GSMNP). A survey of distributions
of the remaining 28 suggests that at least six more are possible from relatively
small families containing uncommonly encountered species. Marske
and Ivie (2003) calculated that 15 families of Coleoptera account for about
80% of total species diversity in the US (Table 1). Based on an analysis of
the distributions of species within these families, we currently estimate that
the total beetle diversity in GSMNP may approach 3000, though estimates as
The Great Smoky Mountains National Park All Taxa Biodiversity Inventory:
A Search for Species in Our Own Backyard
2007 Southeastern Naturalist Special Issue 1:183–192
184 Southeastern Naturalist Special Issue 1
Table 1. Current inventory status of the 15 largest Coleoptera (beetle) families from the US and
Canada in GSMNP (species numbers from Marske and Ivie 2003).
# species # species # species
in US and documented estimated Source
Coleoptera families Canada GSMNP GSMNP authority
Staphylinidae (rove beetles) 4153 248 800 C. Carlton
Curculionidae (weevils) 2919 128 300+ R. Anderson
Carabidae (ground beetles) 2402 286 250 J. Ciegler
Chrysomelidae (leaf beetles) 1869 205 350 E. Riley
Scarabaeidae (scarab beetles) 1700 98 300 M. Paulson
Tenebrionidae (darkling beetles) 1184 59 100 C. Triplehorn
Elateridae (click beetles) 965 99 90 A. Mayor
Cerambycidae (long-horned beetles) 958 159 200 F. Hovore
Buprestidae (metallic wood boring beetles) 762 23 50 C. Bellamy
Melyridae (soft-winged fl ower beetles) 520 6 25 A. Mayor
Dytiscidae (predacious diving beetles) 513 51 45 R. Roughly
Coccinellidae (ladybird beetles) 481 38 50 J. Chapin
Cantharidae (soldier beetles) 473 31 70 S. Kazantsev
Anobiidae (anobiid and spider beetles) 471 12 104 K. Philips
Histeridae (hister beetles) 435 45 65+ A. Tishechkin
Total 19,805 1488 2799+
high as 15,000 have been circulated and our initial estimate was 6000. Our
more conservative figure is based on estimates of species expected for the
15 largest US beetle families (Table 1) in eastern North America plus 20%
to account for species in the remaining families. The currently documented
number of 2131 species demonstrates the incomplete stage of the inventory,
but represents substantial progress from the ≈700 species that were known
from the park six years ago.
In this paper, we review our six-year involvement with the GSMNP
ATBI. We discuss the current status of taxa (especially species new to science)
describe ancillary research and other activities associated with the
project, and outline our plan for completing the GSMNP beetle inventory
during the next three years.
Our basic operational strategy for conducting this inventory has been
to use the taxonomic working group (TWIG; White and Morse 2000).
The Coleoptera TWIG operates as a consortium of professional and avocational
taxonomists, students, and volunteers who contribute different
kinds of talents and expertise towards the common goal of documenting the
beetle fauna of GSMNP. Coleoptera samples are consolidated, sorted, and
processed for distribution to cooperators in the Lousiana State Arthropod
Museum (LSAM) lab. Taxonomic emphasis on particular beetle families
varies according to the resource and time budgets of cooperators and our
ability to sort and deliver these groups for processing. Cooperators are
asked to return identified vouchers to the collection housed at GSMNP and
provide taxonomic and specimen-level data to the LSAM for integration into
2007 C. Carlton and V. Bayless 185
the ongoing inventory checklist. Much taxonomic work is also conducted
by faculty, staff, and students in the LSAM. In addition to families covered
by LSAM specialists, taxa that are not currently being studied by available
specialists and common species that specialists are not enthusiastic about
are identified in-house. The LSAM’s project website (http://entomology.
lsu.edu/lsam/smokybeetles.htm) serves two equally important functions by
consolidating basic species inventory information and disseminating information
about the project to cooperators and the rest of the world.
Data capture is accomplished in the LSAM using Biota (Colwell
2002), database software designed specifically for entomological data
capture. Standard collection and taxonomic data fields are entered and
linked to unique specimen numbers. Periodically all GSMNP Coleoptera
specimen records are exported and transferred to data managers at GSMNP
and Discover Life in America, Inc. (DLIA), where they are imported
to their data management system. DLIA is the non-profit organizational
partner in the GSMNP ATBI and serves as a liaison in coordinating many
aspects of the research.
Coleoptera samples generated by the structured protocol phase of the ATBI
(White and Morse 2000) have contributed the majority of specimens processed
to date through the LSAM. Additional fieldwork has been conducted
mainly through two kinds of activities. Bioblitzes focused on Coleoptera were
conducted during 2001, 2003, 2005, and 2006. The Coleoptera TWIG also
participated in the 2004 Litter Bioblitz. These events provided a format for
specialists, students, and volunteers to collect intensively using a wide variety
of techniques. Annual or semiannual field trips were conducted by the LSAM
lab group to fill seasonal and habitat gaps in sampling coverage.
Educational presentations to students, citizen scientists, and park visitors
are an integral part of the beetle-inventory project. These are organized and
coordinated through DLIA during the bioblitzes and lab field trips. Beginning
in 2007, a series of teacher workshops will be held that will focus on
using beetles as model organisms in primary and secondary science education.
The project provides a living laboratory for students in the LSAM’s
systematics and ecology graduate and undergraduate training program.
When the LSAM became involved in this project during early 2001, approximately
700 species of beetles were authoritatively documented from
GSMNP. This figure was based on two lists, one representing identified
species in the GSMNP collection, and another compiled by avocational
coleopterists Janet Ciegler and Will Merritt, who had conducted extensive
sampling in the park during 1999–2000. Six years of activity by the Coleoptera
TWIG has produced a steadily increasing accumulation of species
(Fig. 1) that currently stands at 2131, based on our database of 16,370
specimens and contributions by cooperators. The current situation can be
understood by focusing on the status of the five largest families and the 42
species that are suspected or known to be undescribed or “new to science.”
186 Southeastern Naturalist Special Issue 1
The largest family of Coleoptera in the US, Staphylinidae (rove beetles
and allies, 4153 species in North America north of Mexico; species numbers
from Marske and Ivie 2003), will almost certainly be the most diverse family in
GSMNP. Currently, we have recorded 248 species and estimate that 800 should
eventually be documented. Of the ten major subfamilies, only one, the Pselaphinae,
is reasonably well inventoried, with 76 species recorded. Six species
of pselaphines (8%) are known to be undescribed, including one new genus,
all of which should be described within the next two years by C. Carlton. The
subfamily Aleocharinae is likely to contain the largest number of undescribed
species of any major taxon based on the large number of undescribed species
in the US as a whole (Newton et al. 2001), but work on this group is hampered
by a severe lack of expertise that was exacerbated by the recent death of the
world's leading expert and Coleoptera TWIG cooperator, Dr. James S. Ashe.
The diversity of moist forest habitats in GSMNP is likely to produce the richest
staphylinid fauna in North America, especially among subfamilies that are
prone to endemism in montane habitats (e.g., Aleocharinae, Omaliinae, and
Pselaphinae). Staphylinids represent ten (24%) of the 42 undescribed species
documented to date (Table 2).
Curculionidae (weevils, 2919 species) constitutes the second largest family
of US beetles. Robert Anderson (Canadian Museum of Nature, Ottawa, ON,
pers. comm.) estimated that 300+ species occur in GSMNP and that 2–5% of
these may be undescribed. Only 123 are currently documented. As an exclusively
phytophagous group, weevils include a great number of pest species,
including the single most important group of forest tree pests, the Scolytinae
(bark and ambrosia beetles), and a large number of non-native “trash” species
(e.g., Carlton and Anderson 2004). Fifty-two species of scolytines are currently
recorded for GSMNP. We estimate that nearly 100 species will eventually be
found, based on figures cited for eastern US (Anderson 2002). This subfamily
is currently under study for the project by Robert Rabaglia (USDA Forest Service,
Carabidae (ground beetles, 2402 North American species) are one of the
better documented families in GSMNP, with 288 species recorded, thanks in
Figure 1. Observed Coleoptera (beetle) species richness for GSMNP project years
2001–2006 and projected richness accumulation through 2009.
2007 C. Carlton and V. Bayless 187
large part to the efforts of J. Ciegler and I.M. Sokolov (LSAM). GSMNP is a
global hotspot for the tribe Cychrini (20 species) and for endemic species in
genera such as Trechus (20 species). One of the more taxonomically intractable
genera, Anillinus, was recently revised. Four of 17 new species were
recorded from GSMNP (Sokolov et al. 2004), and two additional species have
since been found (Sokolov et al., 2007). The exclusively phytophagous leaf
beetles (Chrysomelidae, 1869 North American species) are also reasonably
well inventoried with 202 species recorded as a result of ongoing survey
efforts by E. Riley (Texas A&M University), S. Clark (Brigham Young
University), and others. Riley expects an additional 150 species based on
host-plant distributions. The recently described Psylliodes appalachianus
Konstantinov and Tishechkin (2004) represented the first record of fl ightless,
leaf litter-inhabiting chrysomelids in the US. It is apparently endemic to intermediate
and higher elevations in GSMNP. Most of the species from GSMNP
that are undescribed are members of North American genera that are in need of
modern revisionary treatments. The group also includes a number of invasive
non-native species, and these have been added to the list of unwelcome park
residents (Staines and Staines 2006).
Scarabaeidae. (scarab beetles, 1700 North American species) are surprisingly
poorly sampled in GSMNP, with only 98 species documented.
Additional collecting using light traps will substantially increase this
number. Matt Paulsen (University of Nebraska, Lincoln, NB, pers. comm.)
estimated 250–300 species eventually may be recorded with few or no undescribed
species expected in this well-studied family.
In summary, well-studied families containing large charismatic species
such as the Cerambycidae (long-horned beetles) and Scarabaeidae
(scarabs) have yielded no undescribed species thus far, and none or very
few are expected. Diverse but poorly known families have contributed the
largest number of undescribed species, but taxonomically intractable genera
from otherwise well-studied groups have also contributed to the total.
When undescribed species are found in a genus that is in need of revision
and no specialist is working on that genus, descriptions are delayed. The
specimen will then be noted only as “new species.” An example of this is
in the arcane family Scydmaenidae (antlike stone beetles), with one North
American specialist. Many North American genera are in need of complete
overhaul; therefore, the prospect of seeing names in the foreseeable future
for the seven undescribed species currently listed from GSMNP is unlikely.
Many of the 12 additional species that are identified only to morphospecies
will also turn out to be undescribed (S. O’Keefe, Morehead State University,
Morehead, KY, pers. comm.).
Some comments about our overall approach to this project may serve to
instruct incipient and current ATBI’s at other natural areas in their approach
to conducting inventories of diverse and unevenly studied taxa such as
beetles. The TWIG methodology has served the basic inventory objectives
188 Southeastern Naturalist Special Issue 1
Table 2. Annotated checklist of Coleoptera (beetle) species new to science documented from Great Smoky Mountains National Park, 2001–present.
Taxa N groupA distribution Current status Comments
Carabidae (ground and tiger beetles)
Anillinus cieglerae Sokolov & Carlton 59 Micropredator GSMNPB Described, Sokolov et al. 2007 A. langdoni species
Anillinus langdoni Sokolov & Carlton 98 Micropredator S. Appalachians Described, Sokolov et al. 2004 A. langdoni species
Anillinus loweae Sokolov & Carlton 232 Micropredator S. Appalachians Described, Sokolov et al. 2004
Anillinus moseleyae Sokolov & Carlton 28 Micropredator GSMNP Described, Sokolov et al. 2004
Anillinus murrayae Sokolov & Carlton 11 Micropredator S. Appalachians Described, Sokolov et al. 2004
Anillinus pusillus Sokolov & Carlton 75 Micropredator GSMNP Described, Sokolov et al. 2007 A. langdoni species
Trechus clingmanensis Donabauer 3 Micropredator GSMNP Described, Donabauer 2005a
Trechus pseudonovaculosus Donabauer 2 Micropredator GSMNP Described, Donabauer 2005a
Trechus ramseyensis Donabauer 1 Micropredator GSMNP Described, Donabauer 2005a
Trechus stefanschoedli Donabauer 10 Micropredator GSMNP Described, Donabauer 2005a
Trechus thunderheadensis Donabauer 33 Micropredator GSMNP Described, Donabauer 2005b
Cantharidae (soldier beetles)
Rhagonycha new spp. (at least 2) 5 Predator Unknown Under study, S. Kazantsev Genus in need of revision
Cerylonidae (cerylonid beetles)
Philothermus stephani Gimmel & Slipinski 13 Fungivore S. Appalachians Described, Gimmel and Slipinski 2007
Chrysomelidae (leaf beetles)
Diachus spp. (2) ? Herbivore Unknown Under study, E. Riley
Epitrix sp. ? Herbivore Unknown Under study, E. Riley
Longitarsus spp. (2) ? Herbivore Unknown Under study, E. Riley
Margaridisa new sp. ? Herbivore Unknown Under study, E. Riley
Psylliodes appalachianus Konstantinov 10 Herbivore GSMNP Described, Konstantinov and Tishechkin 2004
Xanthonia sp. ? Herbivore Unknown Under study, E. Riley
2007 C. Carlton and V. Bayless 189
Table 2, continued.
Taxa N groupA distribution Current status Comments
Leiodidae (round fungus and small carrion beetles)
Ptomaphagus merritti Tishechkin 6 Fungivore? S. Appalachians Described, Tishechkin, 2007
Mycetophagidae (hairy fungus beetles)
Triphyllus sp. (possible n. sp., but identity uncertain) 2 Fungivore GSMNP, Under study, R. Leschen First record of genus in
possibly Europe North America
Phalacridae (shining fl ower beetles)
Acylomus new sp. ? Fungivore Eastern US Under study, M. Gimmel and
Scydmaenidae (antlike stone beetles)
Chelonoidum new sp. (det. O'Keefe) 1 Micropredator GSMNP Genus in need of revision
Euconnus new spp. (at least 3, det. S. O'Keefe) 65 Micropredator GSMNP Megadiverse genus in need
Microscydmus new sp. 1 Micropredator GSMNP Genus in need of revision
Stenichmus new sp. 5 Micropredator GSMNP Genus in need of revision
Veraphis new sp. 1 Micropredator GSMNP Genus in need of revision
Staphylinidae (rove beetles)
Arianops new sp. 1 Micropredator GSMNP Under description, C. Carlton
Batrisodes new spp. (at least 2, det. C. Carlton) 13 Micropredator GSMNP Under study, C. Carlton Genus in need of revision
Batrisodes “spretoides” 5 Micropredator GSMNP Under study, C. Carlton O. Park manuscript name
Dasycerus new sp. ? Fungivore GSMNP Under study, V. Gusarov
Geostiba new spp. (possibly several) ? Fungivore GSMNP Under study, V. Gusarov
Gyrophaena new sp. (det. S. Ashe) ? Fungivore GSMNP
New genus, new species 6 Micropredator GSMNP Under study, C. Carlton
Reichenbachia new sp. 2 Micropredator GSMNP Under study, C. Carlton
Rhexius new sp. 5 Micropredator GSMNP Under study, C. Carlton
ASlash indicates functional group of larvae/adult.
BGSMNP = Great Smoky Mountains National Park
190 Southeastern Naturalist Special Issue 1
for Coleoptera reasonably well, but not always in ways that we expected.
The idea of seamless delivery of precisely sorted taxa to interested specialists
for detailed taxonomic work has proven unrealistic. For example, some
malaise-trap samples may contain hundreds of specimens of one species,
and many different traps and trap dates will contain the same species.
Unless the sorter knows how to recognize morphospecies so that they do
not mount all specimens of the same species, then the specialist stands to
receive hundreds of that common species. They often are unable or unwilling
to spend time handling and identifying so many duplicate specimens.
However, because valuable material may lurk at low densities among these
specimens and the distributional and habitat data must be captured, the sorting
can only be accomplished by personnel having at least moderate levels
of taxonomic expertise that can recognize the families and can precisely sort
morphospecies. Staffing labs with individuals at this level of expertise and
dealing with the vast numbers of specimens from mass-collecting methods
have proven to be the most challenging aspects of the project.
No asymptote seems imminent in our observed and projected speciesaccumulation
curve (Fig. 1), and authoritative estimates of potential species
diversity in the top 15 families (Table 1) suggest that a great deal of work is yet
to be done. Interestingly, the actual species numbers for three families, Carabidae
(ground and tiger beetles), Elateridae (click beetles), and Dytiscidae
(predacious diving beetles), have already equaled or slightly surpassed the estimates,
suggesting that either our inventory is complete for these families or,
more likely, our estimate of 3000 beetle species may be too low.
In addition to completing the basic inventory work and shepherding as
many new species through the process of description as possible, several
large-scale products are being prepared for release during the next three
years. Species web pages for the ATBI are being created and posted on the
DLIA website (www.dlia.org) and our LSAM website (http://entomology.
lsu.edu/lsam/smokieschecklist.htm). These include images and distributional,
taxonomic, and life-history information. Posting individual web
pages for all 3000 species is probably unrealistic. A more reasonable goal
is planned whereby at least one species from each genus (estimated at 800)
will be represented. Several researchers have made significant inroads on
webpages in their areas of expertise (e.g., E. Riley [Chrysomelidae, http://
insects.tamu.edu/research/collection/chrysomelidae/] and C. and S. Staines
[aquatic Coleoptera, DLIA website]). A hardcopy annotated checklist and
identification guide to GSMNP Coleoptera is also planned. Contributions
from cooperators and a standard format for chapters will build on the precedent
set by the web-based species pages. The volume’s purpose will be: to
list species that occur in GSMNP; provide notes on their habitats, habits,
and distributions; and to the extent possible, provide users the information
necessary to make identifications.
Funding was provided by an NSF Biotic Surveys and Inventory grant (DEB
0516311 to C.E. Carlton and V.M. Bayless) and mini-grants from DLIA. We thank
2007 C. Carlton and V. Bayless 191
all our TWIG cooperators, Keith Langdon, Becky Nichols, and the inventory and
monitoring staff and rangers at GSMNP. We also thank Jeanie Hilten and the staff
of DLIA, Chuck Parker (US Geological Survey), and the many volunteers, students,
and ATBI colleagues for their support and inspiration. We especially thank Adriean
Mayor, GSMNP Museum Curator, for “ground truthing” many of the older records
and for providing many new ones. This manuscript is approved by the Director, Louisiana
State Agricultural Experiment Station, as manuscript number 07-26-0147.
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