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Comparison of Coleoptera Emergent from Various Decay Classes of Downed Coarse Woody Debris in Great Smoky Mountains Na- Tional Park, USA

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Comparison of Coleoptera Emergent from Various Decay Classes of Downed Coarse Woody Debris in Great Smoky Mountains Na- Tional Park, USA INSECTA A Journal of World Insect Systematics MUNDI 0260 Comparison of Coleoptera emergent from various decay classes of downed coarse woody debris in Great Smoky Mountains Na- tional Park, USA Michael L. Ferro Louisiana State Arthropod Museum, Department of Entomology Louisiana State University Agricultural Center 402 Life Sciences Building Baton Rouge, LA, 70803, U.S.A. [email protected] Matthew L. Gimmel Division of Entomology Department of Ecology & Evolutionary Biology University of Kansas 1501 Crestline Drive, Suite 140 Lawrence, KS, 66045, U.S.A. [email protected] Kyle E. Harms Department of Biological Sciences Louisiana State University 202 Life Sciences Building Baton Rouge, LA, 70803, U.S.A. [email protected] Christopher E. Carlton Louisiana State Arthropod Museum, Department of Entomology Louisiana State University Agricultural Center 402 Life Sciences Building Baton Rouge, LA, 70803, U.S.A. [email protected] Date of Issue: November 30, 2012 CENTER FOR SYSTEMATIC ENTOMOLOGY, INC., Gainesville, FL M. L. Ferro, M. L. Gimmel, K. E. Harms and C. E. Carlton Comparison of Coleoptera emergent from various decay classes of downed coarse woody debris in Great Smoky Mountains National Park, USA Insecta Mundi 0260: 1–80 Published in 2012 by Center for Systematic Entomology, Inc. P. O. Box 141874 Gainesville, FL 32614-1874 USA http://www.centerforsystematicentomology.org/ Insecta Mundi is a journal primarily devoted to insect systematics, but articles can be published on any non-marine arthropod. Topics considered for publication include systematics, taxonomy, nomenclature, checklists, faunal works, and natural history. Insecta Mundi will not consider works in the applied sciences (i.e. medical entomology, pest control research, etc.), and no longer publishes book reviews or editorials. In- secta Mundi publishes original research or discoveries in an inexpensive and timely manner, distributing them free via open access on the internet on the date of publication. Insecta Mundi is referenced or abstracted by several sources including the Zoological Record, CAB Abstracts, etc. Insecta Mundi is published irregularly throughout the year, with completed manuscripts assigned an individual number. Manuscripts must be peer reviewed prior to submission, after which they are reviewed by the editorial board to ensure quality. One author of each submitted manuscript must be a current member of the Center for Systematic Entomology. Managing editor: Paul E. Skelley, e-mail: [email protected] Production editors: Michael C. Thomas, Brian Armitage, Ian Stocks Editorial board: J. H. Frank, M. J. Paulsen Subject editors: G.B. Edwards, J. Eger, A. Rasmussen, F. Shockley, G. Steck, Ian Stocks, A. Van Pelt, J. Zaspel Spanish editors: Julieta Brambila, Angélico Asenjo Printed copies (ISSN 0749-6737) deposited in libraries of: CSIRO, Canberra, ACT, Australia Museu de Zoologia, São Paulo, Brazil Agriculture and Agrifood Canada, Ottawa, ON, Canada The Natural History Museum, London, Great Britain Muzeum i Instytut Zoologiczny PAN, Warsaw, Poland National Taiwan University, Taipei, Taiwan California Academy of Sciences, San Francisco, CA, USA Florida Department of Agriculture and Consumer Services, Gainesville, FL, USA Field Museum of Natural History, Chicago, IL, USA National Museum of Natural History, Smithsonian Institution, Washington, DC, USA Zoological Institute of Russian Academy of Sciences, Saint-Petersburg, Russia Electronic copies (On-Line ISSN 1942-1354, CDROM ISSN 1942-1362) in PDF format: Printed CD mailed to all members at end of year. Florida Center for Library Automation: http://purl.fcla.edu/fcla/insectamundi University of Nebraska-Lincoln, Digital Commons: http://digitalcommons.unl.edu/insectamundi/ Goethe-Universität, Frankfurt am Main: http://edocs.ub.uni-frankfurt.de/volltexte/2010/14363/ Author instructions available on the Insecta Mundi page at: http://www.centerforsystematicentomology.org/insectamundi/ Copyright held by the author(s). This is an open access article distributed under the terms of the Creative Com- mons, Attribution Non-Commercial License, which permits unrestricted non-commercial use, distribution, and reproduction in any medium, provided the original author(s) and source are credited. http://creativecommons.org/ licenses/by-nc/3.0/ 0260: 1-80 2012 Comparison of Coleoptera emergent from various decay classes of downed coarse woody debris in Great Smoky Mountains National Park, USA Michael L. Ferro Louisiana State Arthropod Museum, Department of Entomology Louisiana State University Agricultural Center 402 Life Sciences Building Baton Rouge, LA, 70803, U.S.A. [email protected] Matthew L. Gimmel Division of Entomology Department of Ecology & Evolutionary Biology University of Kansas 1501 Crestline Drive, Suite 140 Lawrence, KS, 66045, U.S.A. [email protected] Kyle E. Harms Department of Biological Sciences Louisiana State University 202 Life Sciences Building Baton Rouge, LA, 70803, U.S.A. [email protected] Christopher E. Carlton Louisiana State Arthropod Museum, Department of Entomology Louisiana State University Agricultural Center 402 Life Sciences Building Baton Rouge, LA, 70803, U.S.A. [email protected] Abstract. Coleoptera species composition and succession in downed woody debris habitats are poorly known in eastern North America. A photoeclector emergence chamber was used to concentrate Coleoptera that emerged from various decay classes of fine and coarse woody debris (FWD and CWD, respectively) collected in primary and secondary forest sites in Great Smoky Mountains National Park, Tennessee, USA. A total of 5673 adult beetle specimens, representing 305 lowest identifiable taxa within 227 genera and 51 families, was collected. One hundred fifteen species (38%) were represented by single individuals. Many more specimens and species were collected from CWD (4129 and 247, respectively) than from FWD (1544 and 162, respectively), but species richness accumulation curves were not significantly different. Many more specimens but an equal number of species were collected from primary forest (3347 and 207, respectively) than from secondary forest (2326 and 207, respectively). Species accu- mulation curves indicated higher richness in secondary forests. Based on a subset of 71 species represented by 10 or more specimens, 27 species were associated with fresh fine woody debris, 11 species with weathered fine woody debris, four with coarse woody debris decay class I, 14 with coarse woody debris decay class II, and eight with coarse woody debris decay class III–IV. Sixteen species were associated with secondary forests, whereas 28 species were associated with primary forests. Coarse woody debris decay class II taken in primary forests had highest absolute species richness with 156 species. In coarse woody debris species overlap decreased with increased difference in decay indicating faunal succession. Published works related to the study of the ecology of downed woody material are briefly summarized. Recommendations on developing a database of legacy trees for future researchers are given. Notes on the biology and photographs of the 71 species represented by 10 or more specimens are given to provide an atlas of eastern U.S. beetle species most commonly encountered in these habitats. “I try all things, I achieve what I can.” ―Herman Melville, Moby-Dick 1 2 • INSECTA MUNDI 0260, November 2012 FERRO ET AL. Introduction Dead trees are unique habitats. No other habitat is 1) a contiguous patch of biological material that; 2) is not actively defended metabolically; 3) is in the presence of plenty of available oxygen; and 4) is persistent for years, decades, or centuries. For the organisms that interact with woody debris, the habitat 1) offers a long term (one to many generations) source of matter and energy; 2) provides mechanical protection from competitors, predators, and parasites; 3) offers patches of reduced environ- mental variability; and is 4) evolutionarily passive (lineages do not evolve in response to the actions of the organisms that inhabit their carcasses). Hamilton (1978) recognized the unique qualities of this habitat and commented on four examples of convergence within the “arbonecrophilic” insect fauna: wing polymorphism; male haploidy; social organization, including subsocial and eusocial; and sexual dimorphism. He postulated that dead wood may have played an important role in the reinvasion of water by terrestrial insects, and may have been the ancestral habitat of progenitors of major insect groups. For example, the evolution of elytra (and therefore Coleoptera) is thought to have resulted from use of the numerous tight spaces provided by dead and dying trees. Support for this is provided by data from the oldest fossil beetles, their wood bor- ings, and the habits of the members of the suborder Archostemata, the most plesiotypic extant beetles (Crowson 1960, 1981; Grimaldi and Engel 2005; Hunt et al. 2007). Attitudes toward dead wood concerning its use as fuel, role in the carbon cycle, and the role it plays in a healthy ecosystem changed greatly at the end of the 1970s and beginning of the 1980s (Speight 1989; Thomas 2002). Studies of dead wood, in particular its role as a reservoir for biodiversity, have recently increased largely due to a desire to use invertebrates as indicators of high quality forests and because many species of conservation concern are also dead wood dependent (Speight 1989; Grove 2002b). Numerous studies of saproxylic organisms, particularly
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