Although the longhorned beetle fauna (Coleoptera: Cerambycidae) is relatively well known across North America, recent efforts have documented substantial increases in longhorned beetle diversity and distribution in Montana (Hart et al. 2013), Nebraska (Spomer 2014), Mississippi (Schiefer 1998, 2001), and Alabama (Holt 2013). Knowledge about the distribution of cerambycid beetles across Tennessee remains limited, and documentation of reporting has been sporadic.

Hoyt Jamerson compiled a list and maps of the longhorned beetle fauna found in Tennessee as part of his Master's thesis at Memphis State University (Jamerson 1973). His examination of published literature, plus field collection efforts and a survey of then-available institutional and some private collections, yielded some 185 species within Tennessee. Data from the thesis were not published, and although it is cited by Linsley & Chemsak (1997), the thesis was miss-catalogued following the integrations of Memphis State University holdings within the University of Memphis Library System (G. P. Barton, personal communication). A portion of Jamerson's specimens have been relocated and integrated into the University of Georgia Collection of Arthropods [UGCA], along with the remainder of the Memphis State University entomology collection. Other collections examined or acknowledged by Jamerson, including specimens from Vanderbilt University, are no longer available or much diminished by damage (e.g., from dermestid beetles and rough handling).

Regardless, since Jamerson's project, considerable monitoring efforts for woodboring beetles have been undertaken across portions of Tennessee. Although recent efforts have been focused on metallic woodboring beetles, bark beetles, and other species of economic interest (Oliver & Mannion 2001; Oliver et al. 2002, 2004; Hansen et al. 2012, 2015; Klingeman et al. 2015), extended-season trapping in the region has yielded many longhorned beetle specimens.

To address existing knowledge gaps, cerambycid specimens were examined at institutional research and teaching collections, museums, and personal collections to assemble available label data and to determine species occurrence and distribution across Tennessee. Label data were examined to inform about seasonal flight activity of adult longhorned beetles and to document, when recorded, the various methods used to collect specimens. When present, relevant notes about adult resources and larval hosts listed for species were compiled.

Documenting the extant cerambycid fauna in Tennessee will elucidate the range of distribution among species and is expected to highlight gaps in collection activity across Tennessee's ecoregions (Fig. 1). Collection data are important for many reasons, including documentation of species diversity, use in assessing environmental and economic impacts of habitat and host plant loss, and monitoring incursion and spread of invasive species. The current specimen summarization effort will update past work to document longhorned beetle species diversity and will help direct future field collection activities.

Supplementary material for this article in Florida Entomologist 100(2) (Jun 2017) is online at  http://purl.fcla.edu/fcla/entomologist/browse. Therefore, the supplementary table is referred to in this article as Suppl. Table 3, and all distributional maps for species are displayed online in supplementary figures (Suppl. Figs. 2–231). Fig. 1 is displayed in color in the online version of the journal.

Materials and Methods

Longhorned beetle specimens were identified using descriptive keys (e.g., Yanega 1996; Lingafelter 2007), and data compiled within this report are derived from these individuals and from examinations of specimens and data from institutional and private collections. Many beetles were also collected in conjunction with a series of seasonlong experimental trials conducted by Tennessee State University affiliated authors and cooperators. A majority of extended seasonal collections that informed this study were made in regions of middle or eastern Tennessee at about weekly intervals from 1 Apr to 28 Aug 2001; 6 Apr to 7 Oct 2002; 5 May to 2 Sep 2003; 12 Apr to 9 Aug 2004; 2 May to 15 Aug 2005; 2 May to 15 Aug 2006; 5 Jun to 21 Jul 2009; 8 Jun to 5 Aug 2010; 17 Jun to 18 Aug 2011; 9 Apr to 10 Sep 2012; and 11 May to 10 Sep 2013.

Specimens were examined in collection depositories [brackets] (Evenhuis 2016, or as acknowledged) at the University of Tennessee Entomology and Plant Pathology Insect Museum [ECUT] and the Great Smoky Mountains National Park [GSNP]. Both collections included results of tree sampling done at multiple times across 2 or more seasons from targeted tree species, for example, tulip poplar (Liriodendron tulipifera L.; Magnoliales: Magnoliaceae) (LaForest et al. 2000), northern red oak (Quercus rubra L.; Fagales: Fagaceae) (Trieff 2002), southern magnolia (Magnolia grandiflora L.; Magnoliales: Magnoliaceae) (Werle 2002), and eastern hemlock (Tsuga canadensis [L.] Carrière; Pinales: Pinaceae) (Buck 2004).

Fig. 1.

Longhorned beetle species tallied within each of the 95 Tennessee counties from collection records compiled for 230 species. Collection distribution is presented across ecoregions occurring within the western, middle, and eastern Grand Divisions of Tennessee (bold black lines). Across the Grand Divisions, county names presented in pale gray text are those from which no longhorned beetle species were collected or reported. Species tallies presented do not include county records reported in Jamerson (1973) that could not be substantiated with a specimen. Roman numerals (west to east) designate the ecoregions of Tennessee, where I corresponds with the Mississippi Alluvial Plain (ecoregion 73), II are the Mississippi Valley Loess Plains (ecoregion 74), III are the Southeastern Plains (ecoregion 65), IV is the Interior Plateau (ecoregion 71), V are the Southwestern Appalachians (ecoregion 68), VI are the Central Appalachians (ecoregion 69), VII are Ridges and Valleys (ecoregion 67), and VIII are the Blue Ridge Mountains (ecoregion 66) (after Griffith et al. 1997). Full descriptions of the Tennessee ecoregions are available at:  https://www.epa.gov/eco-research/ecoregion-download-files-state-region-3.

Additional collections and published records of collections with Tennessee specimen data that were evaluated and integrated into this report include: [CMNH], [CSCA], [CUAC], [CUIC], [ECUT], [EDNC], [GSNP], [FSCA], [LSAM], [MCPM], [MEM], [PERC], [TSRS], [UGCA], [UMMZ], and [UTCI]. [UGCA] also recently had acquired and made available the arthropod collection previously maintained at the Memphis State University (now, University of Memphis). Prionini examined at [MEM] included Tennessee specimens on loan from [AMNH], [CMNH], [CNC], [CSUC], [FSCA], [MCZ], [MSUC], [NYSM], [RAAC], [SCUC], [SEMC], [TAMU], [UAAM], [UDCC], [UKIC], [USNM], and [WIRC] (Evenhuis 2016). Records of observed specimens and published accounts from additional institutional and personal collections that were incorporated included those of: CNTC = Carson—Newman Teaching Collection, DCHS = Davy Crockett High School (Jonesborough, Tennessee), ETSU = East Tennessee State University (Johnson City, Tennessee), FSGA = Forest Service (Dr. Daniel Miller, Athens, Georgia), JMBC = Jason M. Basham, JPBC = Joshua P. Basham, JHGC = Jeffrey P. Huether, NEAC= Nashville Ellington Agriculture Collection, NNYC = Nadeer N. Youssef, TTUC = Tennessee Technological University, WEKC = William E. Klingeman, and WPNC = Warner Parks Nature Center.

Other records that included seasonal activity and county occurrence data were retrieved from published reports (Meyer 1937; Dillon & Dillon 1941, 1947; Linsley 1962a,b, 1963, 1964; Dillon 1956a,b,c; Linsley & Chemsak 1972, 1976, 1984, 1995; TCEIR 1972; McCauley & Eanes 1987; MacRae 2000; Gryzmala 2006; Schiefer & Newell 2010; Tindall et al. 2010; Leavengood & Chapman 2014; Vlasak 2014; Miller et al. 2015).

With the exception of Disteniinae, which is retained within Cerambycidae as a subfamily, nomenclature follows Bezark (2016). Taxonomy is updated to reflect current status, with synonyms used in Jamerson (1973) and more recent publications included after the valid species name. In Table 1, new state records are listed in bold, with specific label and collection data provided. When provided, data were compiled from specimen labels to include collection date, county and locality information, method of collection, and information about habitat, adult resource, or larval plant host. Seasonal adult activities were recorded by range of months, with first and last followed by a numeric superscript indicating week (within month) of first and final collection. For ease of reporting, days 29 to 31 are included within the 4th week period. Observations of species occurrence within any of Tennessee's 8 ecoregions are noted.

The total dataset for species tallies that were tallied in each Tennessee county was analyzed with EstimateS software, using the biascorrected Chao1 (Colwell 2006) that, with 95 iterations, enabled a prediction of the number of species expected within the state and consequently yielding an estimated number of species that remain to be detected.

As with our distribution report on Buprestidae occurring in Tennessee and North Carolina (Klingeman et al. 2015), this report on Tennessee Cerambycidae is presented with inherent bias. Not all portions of the state have been subjected to season-long sampling, and some regions remain largely unexamined. Regardless, this report highlights poorly studied ecoregions across Tennessee and is expected to encourage a more detailed future examination of fauna occurring in these areas.

Table 1.

List of the Cerambycidae of Tennessee with notes on seasonal adult activities and ecoregional distribution(s) (west to east), as noted from collection labels.

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Results

More than 9,918 Tennessee longhorned beetle specimens were assessed, and published literature was reviewed to document 230 species in Tennessee. For some species (e.g., Neandra brunnea [F.]), early or late “outlier” records are noted when specimens were collected across a gap in time that spanned at least 4 wk (Table 1). Total number species occurring in each of Tennessee's 8 ecoregions are reported (Table 2). For each species, available host and habitat data were also compiled from labels and are presented along with managing collection information and noted occurrence within Tennessee counties and ecoregions (Suppl. Table 3; Suppl. Figs. 2–231).

Twenty-seven species are reported as new state records for Tennessee, including Tragosoma harrisii (LeConte), Derobrachus brevicollis Audinet-Serville, Arhopalus foveicollis (Haldeman), Tetropium schwarzianum Casey, Charisalia americana (Haldeman), Grammoptera exigua (Newman), Trigonarthris minnesotana (Casey), Stenocorus ( Stenocorus) schaumii (LeConte), Callidium texanum Schaeffer, Semanotus amethystinus (LeConte), Clytoleptus albofasciatus (Laporte & Gory), Xylotrechus aceris Fisher, Xylotrechus nitidus (Horn), Curius dentatus Newman, Anelaphus moestus moestus (LeConte), Molorchus bimaculatus semiustus (Newman), Obrium rufulum Gahan, Lepturges (Lepturges) pictus (LeConte), Sternidius punctatus (Haldeman), Styloleptus biustus biustus (LeConte), Urgleptes foveatocollis (Hamilton), Urgleptes signatus (LeConte), Oplosia nubila (LeConte), Oberea affinis Leng & Hamilton, Oberea praelonga Casey, Mecas (Mecas) cineracea Casey, and Saperda puncticollis Say. Distribution records for all 230 cerambycid species documented in Tennessee, including potential range additions taken from Jamerson (1973), are plotted by county (Suppl. Figs. 2–231).

Species counts are summed within 85 of the 95 Tennessee counties from which longhorned beetles are reported. Ten Tennessee counties, including Bedford, Bledsoe, Clay, Fentress, Hancock, Marshall, Meigs, Overton, Pickett, and Stewart counties. yielded no taxa among collections and are named in gray shade followed by a “0″ designation. County names are also overlaid upon the ecoregions present among the Grand Divisions of Tennessee, as indicated by bold vertical lines (Fig. 1). Across the top of the map, Roman numeral I corresponds with the Mississippi Alluvial Plain (ecoregion 73) characterized by Northern Mississippi Alluvial Plain habitats, II = Mississippi Valley Loess Plains (ecoregion 74), which includes both Bluff Hills and Loess Plains habitats, III = Southeastern Plains (ecoregion 65), which includes Blackland Prairie, Flatwoods/Alluvial Prairie Margins, Southeastern Plains and Hills, Fall Line Hills, and Transition Hills habitats, IV = Interior Plateau (ecoregion 71), which includes Western Pennyroyal Karst, Western Highland Rim, Eastern Highland Rim, Outer Nashville Basin, and Inner Nashville Basin habitats, V = Southwestern Appalachians (ecoregion 68), which includes Cumberland Plateau, Sequatchie Valley, and Plateau Escarpment habitats, VI = Central Appalachians (ecoregion 69), characterized by Cumberland Mountain habitats, VII = Ridge and Valley (ecoregion 67), which includes Southern Limestone/Dolomite Valleys and Low Rolling Hills, Southern Shale Valleys, Southern Sandstone Ridges, and Southern Dissected Ridges and Knobs habitats, and VIII = Blue Ridge Mountains (ecoregion 66), which includes Southern Igneous Ridges and Mountains, Southern Sedimentary Ridge, Limestone Valleys and Coves, and Southern Metasedimentary Mountains habitats (after Griffith et al. 1997). Counts of species collected within each ecoregion, along with expanded descriptions of the Tennessee ecoregional typography and their associated flora are presented (Table 2) with additional details available at:  https://archive.epa.gov/wed/ecoregions/web/html/tn_eco.html.

Table 2.

Ecoregional distribution of Cerambycidae species tallied within each of the 8 ecoregions occurring across Tennessee.

As with Buprestidae in Tennessee (Klingeman et al. 2015), the areas of most intensive collection activity were focused in eastern Tennessee around the University of Tennessee Knoxville campus and the Great Smoky Mountains National Park. In middle Tennessee, counties around the Otis L. Floyd Research Center in McMinnville were well documented. Remnants of Jamerson's thesis collection (1973) confirmed the presence of many species in the southwestern corner of Tennessee. Likewise, collections made by staff at [MEM] and in conjunction with Cooperative Agricultural Pest Surveys and Invasive Species collections, with wood-associated beetles curated by staff at [PERC], yielded numerous specimens in middle and western Tennessee, along Tennessee portions of the Natchez Trace National Scenic Trail and among cedar glades habitats within the Interior Plateau ecoregion of middle Tennessee.

The Chao1 estimator predicted that 241 (± 6.7 SD) cerambycid beetle species are expected to occur in Tennessee, with an associated confidence interval of 234 to 264 species that may be found within the state.

Discussion

Although several cerambycid species periodically can cause economic injury to forest tree species (e.g., Enaphalodes atomarius [Drury], E. cortiphagus [Craighead], E. rufulus [Haldeman]), herbaceous plants, and row crops (e.g., Dectes sp. on soybean, Glycine max [L.] Merrill, Fabales: Fabaceae), none among the 230 documented longhorned beetle species in Tennessee is considered an exotic invasive species.

The limited collections of longhorned beetle species occurring across the Mississippi Alluvial and Valley Loess Plains, Southeastern Plains, western portions of the Interior Plateau ecoregions, and Central Appalachian region indicates that future collection activity across these Tennessee ecoregions would be expected to more clearly define species intrastate distributions and potential range constraints across the cerambycid fauna of Tennessee (Table 2; Suppl. Figs. 2–231).

The Chao1 estimator suggests that there are likely to be about 11 more longhorned beetle species that occur in Tennessee. Among the 185 longhorned beetle species reported in Jamerson's (1973) thesis are several for which no physical specimen could be located and no associated collection information was provided. Among these nonvalidated accounts are 10 species that may yet be documented in Tennessee, including Parandra polita Say, Leptura abdominalis (Haldeman), L. plebeja Randall, Lycochoriolaus lateralis (Olivier), Strangalia sexnotata Haldeman, Megacyllene decora (Olivier), Enaphalodes hispicornis (L.), Batyle ignicollis australis Linsley, Oberea gracilis (F.), and Saperda obliqua Say.

Deciduous plants are important reproductive hosts for several species that may yet be collected in Tennessee. Parandra polita are attracted to lights (Holt 2013), and larvae of this species develop within decaying heartwood of, for example, Carya (Fagales: Juglandaceae), Fagus (Fagales: Fagaceae), and Liriodendron species (Linsley & Chemsak 1997), which occur across Tennessee. Glycobius speciosus (Say) larvae develop within Acer saccharum Marshall (Sapindales: Sapindaceae), and this beetle species may have been collected in Tennessee in McMinn County, although Holland (2009) does not list the collection that would have contained the record. In Alabama, Holt (2013) reported G. speciosus from Jackson County in ecoregion 68. Oak species are also larval host plants for E. hispicornis, which has been collected in Alabama from Lawrence and Madison counties (ecoregions 71 and 68, respectively) (Holt 2013). Oberea gracilis larvae also develop in seedlings of Quercus alba L. and Quercus falcata Michaux (Fagales: Fagaceae), which occur across Tennessee, yet reported O. gracilis collections were restricted to southern counties in Mississippi and Alabama (Schiefer 1998; Holt 2013). Quercus, Amelanchier (Rosales: Rosaceae), and Castanea (Fagales: Fagaceae) species are larval host plants of Hebestola nebulosa Haldeman. Liriodendron and Nyssa (Cornales: Cornaceae) species are developmental hosts for Aegomorphus morrisii (Uhler) (Linsley & Chemsak 1997), yet A. morrisii adults can also be attracted to a blacklight (Shieffer 1998). Alnus serrulata (Aiton) Willd. (Fagales: Betulaceae), Betula species (Fagales: Betulaceae), and Corylopsis species (Saxifragales: Hamamelidacea) are larval host plants used by S. obliqua (Lingafelter 2007). All 3 of these cerambycid species have been collected in Mississippi in ecoregion 65 (Schiefer 1998), which extends north into Tennessee. Amorpha fruticosa L. (Fabales: Fabaceae), which is the developmental host plant used by M. decora (Linsley & Chemsak 1997), can be found in ecoregions occurring across Tennessee but is less common in ecoregions 69, 66, and 65 (Ma 2016).

Coniferous plants are also used as reproductive hosts with L. abdominalis relying on Taxodium distichum (L.) Richard (Pinales: Cupressaceae) and Juniperus species (Pinales: Cupressaceae), whereas L. plebeja larvae develop within wood of Picea and Pinus species (all Pinales: Pinaceae) (Linsley & Chemsak 1997). Adults of L. plebeja have been collected on Spiraea alba var. latifolia (Aiton) Boivin (Rosales: Rosaceae) flowers and in blue, white, and yellow pan traps in ecoregion 66 in Polk and Henderson counties, in North Carolina (Campbell et al. 2007). Batyle ignicollis australis and Scaphinus muticus (F.), which both can develop within Pinus species, and within herbaceous Bidens species (Asterales: Asteraceae) and Hypericum species (Malpighiales: Hypericaceae), have been collected from southern Mississippi and Alabama (Schiefer 1998; Holt 2013).

Reproductive hosts are not documented for L. lateralis and S. sexnotata, yet adults of these species have been taken on Callicarpa americana L. (Lamiales: Lamiaceae), and on Daucus carota L. (Apiales: Apiaceae) and Bidens aristosa (Michaux) Britton (Asterales: Asteraceae), respectively, in Mississippi ecoregions 74 and 65 (Lago & Mann 1987; Schiefer 1998), as well as in Lauderdale county, Alabama, in ecoregion 71 (Holt 2013).