022 426C Carlton

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

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 Arthro- pod 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 (min- ute 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 mass-collecting methods.

Introduction 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 rela- tively 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

1Louisiana State Arthropod Museum, Department of Entomology, Louisiana State University Agricultural Center, Baton Rouge, LA 70803. *Corresponding author - [email protected].

183 184 Southeastern Naturalist Special Issue 1 high as 15,000 have been circulated and our initial estimate was 6000. Our more conservative ﬁ gure 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.

Methods 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 avo- cational 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 identiﬁ ed vouchers to the collection housed at GSMNP and provide taxonomic and specimen-level data to the LSAM for integration into

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+

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 identifi ed in-house. The LSAM’s project website (http://entomology.lsu.edu/ lsam/smokybeetles.htm) serves two equally important functions by consoli- dating 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 co- ordinating 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 fi eldwork 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 fi eld trips were conducted by the LSAM lab group to fi ll 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 fi eld trips. Begin- ning in 2007, a series of teacher workshops will be held that will focus on using beetles as model organisms in primary and secondary science educa- tion. The project provides a living laboratory for students in the LSAM’s systematics and ecology graduate and undergraduate training program.

Results When the LSAM became involved in this project during early 2001, approximately 700 species of beetles were authoritatively documented from GSMNP. This fi gure was based on two lists, one representing identifi ed species in the GSMNP collection, and another compiled by avocation- al coleopterists Janet Ciegler and Will Merritt, who had conducted extensive sampling in the park during 1999–2000. Six years of activity by the Coleop- tera 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 fi ve 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 Pse- laphinae, 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 rich- est 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 fi gures cited for eastern US (Anderson 2002). This subfamily is currently under study for the project by Robert Rabaglia (USDA Forest Ser- vice, Arlington, VA). 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 fi rst record of fl ightless, leaf litter-inhabiting chrysomelids in the US. It is apparently endemic to inter- mediate 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 sur- prisingly 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 identifi ed only to morphospecies will also turn out to be undescribed (S. O’Keefe, Morehead State University, Morehead, KY, pers. comm.).

Discussion 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 for 188 Southeastern Naturalist Special Issue 1

species

2001–present.

Comments

A. langdoni

A. langdoni complex

, E. Riley

d, Sokolov et al. 2007

d, Sokolov et al. 2004

d, Sokolov et al. 2004 d, Sokolov et al. 2004

d, Sokolov et al. 2004 d, Sokolov et al. 2007 d, Donabauer 2005a

d, Donabauer 2005a d, Donabauer 2005a d, Donabauer 2005a d, Donabauer 2005b

d, Gimmel and Slipinski 2007

Describe complex

Describe Describe Describe

Describe Describe

Describe Describe Describe

Describe

distribution Current status distribution Current

/herbivore

? Herbivore Unknown study Under ? Herbivore Unknown E. Riley Under study, ? Herbivore Unknown study Under ? Herbivore Unknown E. Riley Under study,

? Herbivore Unknown E. Riley Under study,

Functional Known

3 Micropredator GSMNP GSMNP 3 Micropredator GSMNP 2 Micropredator 1 Micropredator GSMNP GSMNP 1 Micropredator

5 Predator Unknown S. Kazantsev Under study, Genus in need of revision

N group N

98 Micropredator S. Appalachians S. 98 Micropredator

28 Micropredator GSMNP GSMNP 28 Micropredator

10 Micropredator GSMNP GSMNP 10 Micropredator

10 Herbivore GSMNP 200 Tishechkin Described, Konstantinov and

Donabauer

Donabauer 33 Micropredator GSMNP GSMNP Micropredator Donabauer 33

Konstantinov

Donabauer

Gimmel & Slipinski 13 Fungivore Appalachians S.

Donabauer

Sokolov & Carlton

Sokolov & Carlton 11 Micropredator Appalachians S.

Sokolov & Carlton 59 Micropredator GSMNP

Sokolov & Carlton

Sokolov & Carlton 75 Micropredator GSMNP

Sokolov & Carlton 232 Micropredator Appalachians S.

stephani

spp. (2)

new sp.

new spp. (at least 2)

appalachianus

sp.

spp. (2)

sp.

Anillinus langdoni

Anillinus loweae

Anillinus moseleyae Anillinus murrayae

Trechus clingmanensis Trechus

Rhagonycha

Diachus

Epitrix

Longitarsus Margaridisa Psylliodes Xanthonia

Table 2. Annotated checklist of Coleoptera (beetle) species new to science documented from Great Smoky Mountains National Park, 2. Table

Taxa Taxa Carabidae (ground and tiger beetles) Anillinus cieglerae

Anillinus pusillus

pseudonovaculosus Trechus Trechus ramseyensis Trechus Trechus stefanschoedli Trechus Trechus thunderheadensis thunderheadensis Trechus Cantharidae (soldier beetles)

Cerylonidae (cerylonid beetles) Philothermus Chrysomelidae (leaf beetles)

Tishechkin &

2007 C. Carlton and V. Bayless 189 W. Steiner W. distribution Current status distribution Current Comments A

Tishechkin 6 Fungivore? S. Appalachians Described, Tishechkin, 2007 Tishechkin, Described, Appalachians S. Fungivore? 6 Tishechkin ower beetles) ower ﬂ new sp. 2 Micropredator GSMNP C. Carlton Under study, new sp. 1 Micropredator GSMNP Genus in need of revision new sp. (det. O'Keefe) 1 Micropredator GSMNP Genus in need of revision new sp. (det. S. Ashe) new sp. (det. S. ? Fungivore GSMNP new sp. 5 Micropredator GSMNP Genus in need of revision new spp. (at least 2, det. C. Carlton) “spretoides” 13 Micropredator GSMNP C. Carlton Under study, Genus in need of revision 5 Micropredator GSMNP C. Carlton Under study, O. Park manuscript name new sp. ? Fungivore GSMNP Gusarov V. Under study, new spp. (at least 3, det. S. O'Keefe) 65 Micropredator GSMNP Megadiverse genus in need sp. (possible n. sp., but identity uncertain) 2 Fungivore GSMNP, R. Leschen Under study, First record of genus in new sp. 1 Micropredator GSMNP Under description, C. Carlton new sp. ? Fungivore Eastern US M. Gimmel and Under study, new spp. (possibly several) ? Fungivore GSMNP Gusarov V. Under study, new sp. 1 Micropredator GSMNP Genus in need of revision new sp. 5 Micropredator GSMNP C. Carlton Under study, Reichenbachia Rhexius Gyrophaena Euconnus Geostiba Ptomaphagus merritti Triphyllus Chelonoidum Microscydmus Stenichmus Batrisodes Batrisodes Dasycerus Veraphis Slash indicates functional group of larvae/adult. GSMNP = Great Smoky Mountains National Park GSMNP

New genus, new species 6 Micropredator GSMNP C. Carlton Under study, Leiodidae (round fungus and small carrion beetles) Scydmaenidae (antlike stone beetles)

Functional Known Functional group 2, continued. Table N Taxa Phalacridae (shining A possibly Europe America North of revision Acylomus Arianops Mycetophagidae (hairy fungus beetles) B Staphylinidae (rove beetles)

190 Southeastern Naturalist Special Issue 1 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 mal- aise-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. Staffi ng 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 species- accumulation 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, Carabi- dae (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 shepherd- ing 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 signifi cant 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 identifi cation guide to GSMNP Coleoptera is also planned. Contributions from cooperators and a standard format for chapters will build on the prec- edent 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 identifi cations.

Acknowledgments 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, Loui- siana State Agricultural Experiment Station, as manuscript number 07-26-0147.

Literature Cited Arnett, R.H., Jr., and M.C. Thomas (Eds.). 2001. American Beetles, Vol. 1: Arch- ostemata, Myxophaga, Adephaga, Polyphaga: Staphyliniformia. CRC Press, Boca Raton, FL. 443 pp. Arnett, R.H., Jr., M.C. Thomas, P.E. Skelley and J.H. Frank (Eds.). 2002. American Beetles, Vol. 2: Polyphaga: Scarabaeoidea through Curculionoidea. CRC Press, Boca Raton, FL. 861 pp. Anderson, R.S. 2002. 131. Curculionidae Latreille 1802 (R.S. Anderson) Pp. 722–815, In R.H. Arnett, Jr., M.C. Thomas, P.E. Skelley, and J.H. Frank (Eds.). American Beetles, Vol. 2: Polyphaga: Scarabaeoidea through Curculionoidea CRC Press, Boca Raton, FL. 861 pp. Carlton, C.E., and R.S. Anderson. 2004. Occurrence of the introduced weevil Myosides seriehispidus Roelofs in Great Smoky Mountains National Park (Cole- optera: Curculionidae). Coleopterists Bulletin 58:343. Colwell, R.K. 2002. BIOTA: The biodiversity database manager. Version 1.6.1. Sinauer Associates, Sunderland, MA. Donabauer, M. 2005a. New species of the Trechus (Microtrechus) uncifer-group from the southern Appalachians (Coleoptera: Carabidae: Trechinae). Zeitschrift der Arbeitsgemeinschaft Österreichischer Entomologen 57:51–62. Donabauer, M. 2005b. New species and subspecies of the Trechus (Microtrechus) nebulosus-group from the southern Appalachians (Coleoptera: Carabidae: Tre- chinae). Zeitschrift der Arbeitsgemeinschaft Österreichischer Entomologen 57: 65–92. Gimmel, M.L., and A. Slipinski. 2007. A new species of the genus Philothermus Aubé (Coleoptera: Cerylonidae) from Great Smoky Mountains National Park, USA. Zootaxa 1390:17–20. Konstantinov, A., and A. Tishechkin. 2004. The ﬁ rst nearctic leaf-litter ﬂ ea beetle (Coleoptera: Chrysomelidae) from the Great Smoky Mountains National Park. Coleopterists Bulletin 58:71–76. Marske, K.A., and M.A. Ivie. 2003. Beetle Fauna of the United States and Canada. Coleopterists Bulletin 57:495–503. Newton, A.F., Jr., M.K. Thayer, J.S. Ashe, and D.S. Chandler. 2001. 22. Staphylini- dae Latreille, 1802. Pp. 272–418, In R.H. Arnett, Jr. and M.C. Thomas (Eds.). American Beetles, Vol. 1: Archostemata, Myxophaga, Adephaga, Polyphaga: Staphyliniformia. CRC Press, Boca Raton, FL. 443 pp. Sokolov, I.M., C.E. Carlton, and J.F. Cornell. 2004. Review of Anillinus with descriptions of 13 new species and a key to soil and litter species (Coleoptera: Trechinae: Bembidiini). Coleopterists Bulletin 58:185–233. 192 Southeastern Naturalist Special Issue 1 Sokolov, I.M., Y.Y. Sokolova, and C.E. Carlton. 2007. New species of Anillinus Casey (Carabidae: Trechinae: Bembidiini) from Great Smoky Mountains Na- tional Park, USA and phylogeography of the A. langdoni species group. Zootaxa 1542: 1-20 Staines, C.L., and S.L. Staines. 2006. Additional eastern US records of the introduced leaf beetle Chrysolina quadrigemina (Suffrain) (Coleoptera: Chrysomelidae). Coleopterists Bulletin 60:72. Tishechkin, A.K. 2007. A new species of Ptomaphagus (Appadelopsis) (Coleop- tera: Leiodidae) from Great Smoky Mountains National Park, USA. Zootaxa 1478: 61–64 White, P., and J. Morse. 2000. The Science Plan for the All Taxa Biodiversity Inven- tory in Great Smoky Mountain National Park. 15 pp. Available online at http: //www.discoverlifeinamerica.org/. Accessed 1 June 2007.