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Annotated Checklist of Some Fungivorous Beetles (Coleoptera: Anamorphidae, Biphyllidae, Derodontidae, Endomychidae, Erotylidae, and Tetratomidae) of the George Washington Memorial Parkway PDF

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Banisteria, Number 50, pages 21-28 © 2018 Virginia Natural History Society Annotated Checklist of Some Fungivorous Beetles (Coleoptera: Anamorphidae, Biphyllidae, Derodontidae, Endomychidae, Erotylidae, and Tetratomidae) of the George Washington Memorial Parkway Brent W. Steury U.S. National Park Service 700 George Washington Memorial Parkway Turkey Run Park Headquarters McLean, Virginia 22101 ABSTRACT Seven collection methods were utilized at a national park site in northern Virginia to capture fungivorous beetles in the families Anamorphidae, Biphyllidae, Derodontidae, Endomychidae, Erotylidae, and Tetratomidae. Forty species and one subspecies were documented, including five new state records (Hallomenus scapularis Melsheimer, Microsternus ulkei [Crotch], Triplax frontalis Horn, Tritoma erythrocephala Lacordaire, and Tritoma mimetica [Crotch]). More than half of the Commonwealth’s expected fauna was documented from the park. Malaise traps were the most productive capture method. Periods of adult activity, based on dates of capture, are given for each species. Relative abundance is noted for each species based on the number of captures. Keywords: Arlington County, Fairfax County, fungus beetles, national park, new state records, Virginia. INTRODUCTION Arlington counties). Park sites that received inventory effort included: in Fairfax County, Dyke Marsh Wildlife Fungi and Coleoptera are among the most diverse Preserve, Fort Marcy, Great Falls Park, Fittle Hunting heterotrophic organisms in the world with an equally Creek, and Turkey Run Park, and in Arlington County, diverse array of commensal and mutualistic fungus- Arlington Woods (at Arlington House). This area covers beetle relations as well as combative interactions approximately 927 ha. A map of these sites is provided including entomopathogenic fungi (Kukor & Martin, in Steury (2011). Great Falls and Turkey Run parks and 1987) and fungivorous beetles (Schigel, 2012). Fort Marcy fall within the Piedmont physiographic Fungivory is characteristic of many families of province while all other collection sites are on the Coleoptera. The fungal mycelia are most frequently Coastal Plain. Most sites are situated along the shore of consumed by beetles together with the woody substrata the Potomac River, and Great Falls and Turkey Run (Schigel, 2012); however, fruit bodies and spores are Parks border the Potomac River Gorge, an area with a also consumed (Schigel, 2008). Fungivorous beetles long history of biodiversity studies (Brown, 2008). Most are not always easy to find in nature, and collecting of the study sites are dominated by maturing, second them requires an arsenal of methods for efficient growth, primarily upland, deciduous woodlands with a documentation (Schigel, 2008). This study utilized seven band of floodplain forest along the Potomac River. More collection methods to assess the fauna of six fungivorous open, herbaceous dominated habitats can be found along beetle families from a national park site in northern the river shores and in the marsh habitats at Dyke Marsh. Virginia. The vascular flora of the GWMP is diverse, with more than 1,313 taxa recorded, 1,020 from Great Falls Park STUDY SITES alone (Steury et al., 2008; Steury, 2011). The study sites include lands managed by the MATERIAFS AND METHODS National Park Service as units of the George Washington Memorial Parkway (GWMP) in Virginia (Fairfax and Specimens were collected during a 19-year period 22 BANISTERIA NO. 50, 2018 (1998-2017) using a variety of sporadic survey efforts the George Washington Memorial Parkway (GWMP), targeting arthropods, including: Malaise traps, Lindgren Turkey Run Park Headquarters in McLean, Virginia. funnels, blacklight (UV) bucket traps, blacklight shone on sheets, leaf litter samples processed in Berlese RESULTS AND DISCUSSION funnels, beating sheets, and hand picking (including examination of fungi in the field). Six Townes style Forty species and one subspecies were documented Malaise traps (Townes, 1962) were set at Dyke Marsh, from GWMP in six families of fungivorous beetles April 1998-December 1999, three each at Great Falls and (Anamorphidae, Biphyllidae, Derodontidae, Endo- Turkey Run parks (March 2006-November 2009), and mychidae, Erotylidae, and Tetratomidae). This is more four at Little Hunting Creek (March-November 2017). than half of the expected fauna of Virginia (see list of Traps at Dyke Marsh were set each year in the same species for number of species expected in Virginia from locations in open, tidal, freshwater marsh dominated by each family), emphasizing the importance of parks as Typha angustifolia L., floodplain forest dominated havens for biodiversity near urban areas. Management by red and silver maple (Acer rubrum L. and A. decisions to not remove, or chip, downed woody debris, saccharinum L.) and tulip poplar (Liriodendron and to leave standing, non-hazardous, dead trees has tulipifera L.), and at the marsh/forest ecotone. In Great undoubtedly provided an abundance of substrates for Falls Park, a trap was set in each of three habitats: quarry fungal growth, and food and shelter for beetles that feed site (dry, upland, mixed deciduous/coniferous forest), on these fungi. Recently, the amount of standing and swamp (dominated by red maple), and floodplain forest downed dead wood in GWMP has greatly increased due (dominated by oaks [Quercus sp.], and tulip poplar). In to the death of large ash trees (Fraxinus americana L., Turkey Run Park, one trap was set in upland forest F. pennsylvanica Marshall, and F. profunda [Bush] dominated by oaks and tulip poplar and two traps in Bush) infected by emerald ash borer (Agrilus floodplain forest along the Potomac River (dominated planipennis Fairmaire), a non-native buprestid beetle. by oaks, basswood [Tilia americana L.], and sycamore Although this loss is detrimental to arthropods that feed [Platanus occidentalis L.]). At Little Hunting Creek, exclusively on ash (Gandhi & Herms, 2010), it will four traps were set in upland forest dominated by an undoubtedly increase habitat and food sources for fungi, ericaceous understory and a canopy of oaks, hickory and the beetles associated with them. (Carya sp.), American beech (Fagus grandifolia Ehrh.), Five species (Hallomenus scapularis, Microsternus and some Virginia pine (Pinus virginiana Mill.). ulkei, Triplax frontalis, Tritoma erythrocephala, and Additional collections of species from these six Tritoma mimetica) are documented in the literature for families of beetles were also made by sporadically using the first time from Virginia. Thirty-four species and one other collecting methods, including running pitfall traps subspecies documented from Great Falls and Turkey set at Dyke Marsh (five years) and at Little Hunting Run parks are recorded for the first time from the Creek and Great Falls and Turkey Run parks (three Potomac River Gorge (Brown, 2008). Only three species years); Lindgren funnel and blacklight (UV) bucket traps (Triplax flavicollis, Tritoma biguttata biguttata, and set at Dyke Marsh, Great Falls Park, Little Hunting Tritoma burneralis) were found to be abundant within the Creek, and Turkey Run Park (two years); blacklight study area. In comparison, the other 38 taxa were rarely shone on sheets at Great Falls and Turkey Run parks encountered despite 19 years of survey effort. Great Falls (three years); leaf litter from Arlington Woods, Dyke Park was the most species-rich site with 28 species, Marsh, Fort Marcy, Great Falls Park, and Turkey Run followed by Turkey Run Park (27 taxa), Little Hunting Park, processed in Berlese funnels (two years); beating Creek (14), and Dyke Marsh (12). Malaise traps were the sheets (three years); and collecting by hand (including most successful capture method (37 taxa), followed by examination of fungi in the field) at all sites, over seven hand picking (12), Berlese funnels (6), Lindgren funnels years. Locations, habitat descriptions, and collection (4), and UV bucket traps (3). methods are summarized in Table 1. Collectors included C. Acosta, E. Barrows, J. Brown, C. Davis, A. Evans, J. LIST OF SPECIES Fisher, S. Lingafelter, D. Mead, E. Oberg, M. Skvarla, D. Smith, W. Steiner, B. Steury, J. Swearingen, and C. The number of specimens in the collection at GWMP Wirth. State record determinations are based on reviews is indicated in parentheses after each taxon. Taxa too of Boyle (1956), Ciegler (2014b), Downie & Arnett common for all specimens to be kept are listed as (1996), Evans (2008), Lawrence (1989), and Shockley et abundant. Sites, habitats, and collection methods are al. (1999). Specimens were determined by Arthur V. given following the abbreviations listed in Table 1. Other Evans or Brent W. Steury. Specimens were pinned, locations and habitats within sites are indicated when labeled, and deposited in the collections maintained at necessary. The periods of adult activity are based on STEURY: FUNGIVOROUS BEETLES 23 Table 1. Summary of locations, latitude and longitude, habitats sampled, and trap types used during this study. Additionally, all sites were sampled by hand picking, including examination of fungi in the field (hp) and beating sheets (bs). Latitude and Location Habitats Sampled Trap Types Longitude Arlington County Arlington Woods (AW) N 38.883, W-77.074 Upland, deciduous forest (uf) Berlese funnels (bf) Fairfax County Berlese funnels; blacklight (UV) Tidal, freshwater marsh (tm); Dyke Marsh Wildlife bucket traps (uv); Lindgren N 38 .772, W -77.050 floodplain, deciduous forest (ff); Preserve (DM) funnels (If); pitfall traps (pt); marsh/forest ecotone (ec) Townes style Malaise traps (mt) Fort Marcy (FM) N 38.937, W-77.125 Upland, deciduous forest Berlese funnels Upland, mixed deciduous/ Berlese funnels; blacklight shone coniferous forest (qu); deciduous on sheets (bl); blacklight bucket Great Falls Park (GF) N 38.985, W -77.246 swamp (sf); floodplain, deciduous traps; Lindgren funnels; pitfall forest traps; Townes style Malaise traps Blacklight bucket traps; Little Hunting Creek Upland, deciduous forest with N 38.717, W-77.078 Lindgren funnels; pitfall traps; (LH) some pine (dp) Townes style Malaise traps Berlese funnels; blacklight shone Upland, deciduous forest; on sheets; blacklight bucket traps; Turkey Run Park (TR) N 38.965, W-77.156 floodplain, deciduous forest Lindgren funnels; pitfall traps; Townes style Malaise traps dates when live individuals have been documented in the Biphyllidae (False Skin Beetles) park. Dates separated by a hyphen indicate that the taxon was documented on at least one day during each month Two species occur in eastern North America, both of within this continuum of months, whereas dates which have been documented in Virginia. One of these separated by a comma represent individual observation species was found in GWMP. These beetles feed on dates. For traps set over multiple weeks, the first day fungal spores (Downie & Arnett, 1996). of the set is used as the earliest date and the last day of the set as the latest date. Taxa newly recorded Diplocoelus brunneus LeConte - (1); LH dp; 19 Sep-10 for the Potomac River Gorge are marked by an asterisk. Oct; mt. Anamorphidae (False Handsome Fungus Beetles) Derodontidae (Tooth-necked Fungus Beetles) This family was split from Endomychidae by Four species in two genera are expected in Virginia, Robertson et al. (2015). Two species have been one of which, Laricobius nigrinus Fender, has been documented from Virginia (Shockley et al., 1999), one introduced as a biological control agent for the hemlock of which was found in GWMP. Species in this family woolly adelgid (Adelges tsugae [Annand]). Laricobius are generally quite small, some less than 1 mm, and thus rubidus LeConte reaches its southern known limit easily overlooked. in Washington, DC, but has not yet been recorded from Virginia (Lawrence, 1989). Laricobius are not *Clemmus minor (Crotch) - (1); TR ff; 18 Aug-4 Sep; fungivores, but instead feed on adelgids (Hemiptera: mt. 24 BANISTERIA NO. 50, 2018 Adelgidae) that feed on conifers. Derodontus feed on fruiting bodies of Basidiomycotic fungi. Two species are documented from GWMP. * Derodontus esotericus Lawrence - (26); DM ec, GF sf, TR ff; 22 May-1 Jul, 19 Sep-5 Dec; mt. *Derodontus maculatus (Melsheimer) (Fig. 1) - (4); GF sf, TR ff; 22 Oct-17 Nov; mt. Always captured in association with D. esotericus. Endomychidae (Handsome Fungus Beetles) Ten species have been documented in Virginia (Shockley et al., 1999), of which seven are recorded from GWMP. Holoparamecus contains five species, three of which are cosmopolitan, but none has been found in Virginia. One of these, H. caularum Aube, has been documented only in the eastern United States from Florida and Michigan. The other two have been recorded near Virginia, H. depressus Curtis (Georgia and New York) and H. singularis (Beck) (New York). A fourth species, H. ragusae Reitter, occurs in Pennsylvania (Shockley et al., 1999). *Aphorista vittata (Fabricius) - (3); GF sf and rocky outcrop above Sandy Landing, TR Leiter Mansion ruins; 15 Apr-10 May, 16-30 Jul; bf, hp, mt. *Endomychus biguttatus Say - (16); DM ec, GF sf, TR uf and ff; 1 May-21 Oct, 15 Dec; hp (under bark of large fallen Liriodendron tulipifera L. [Magnoliales: Magnoliaceae], on fungus), mt. *Mycetina perpulchra (Newman) - (7); GF sf, LH dp, TR ff; 23 Apr-26 Jul; mt. *Phymaphora pulchella Newman - (2); GF sf; 18 Mar- 30 Apr; mt. *Rhanidea unicolor (Ziegler) - (8); AW uf, DM ec, GF qu and Difficult Run, LH dp; 10 Apr-18 May; bf, If, pf, mt. Fig. 1. Derodontus maculatus (Melsheimer); Collection data: Virginia, Fairfax Co., Great Falls Park, swamp, Malaise trap, *Stenotarsus blatchleyi Walton - (12); DM ff, GF sf, LH 22 October - 11 November 2008, B. Steury & D. Smith. Top, dp, TR ff; 19 Jun-14 Aug; mt. dorsal habitus. Bottom, close-up of the head and pronotum of the same specimen. This species was not included for the northeastern United States by Downie & Arnett (1996.). It reaches its northern extreme in Virginia and has not been Stenotarsus hispidus (Herbst) - (1); LH dp; 20-30 Jun; documented in North Carolina (Shockley et al., 1999). mt. STEURY: FUNGIVOROUS BEETLES 25 Erotylidae (Pleasing Fungus Beetles) *Triplax frontalis Horn - (3); AW uf, GF qu; 10-30 Apr; bf, mt. NEW STATE RECORD. There are 57 potential species in South Carolina (Ciegler, 2014a). Downie & Arnett (1996) report 36 This beetle occurs from Nebraska and Texas species in northeastern North America and Evans (2014) eastward to Georgia and Pennsylvania (Boyle, 1956; reports 40 in the eastern United States. Twenty species Downie & Arnett, 1996). and one subspecies have been documented from GWMP. *Triplax thoracica (Say) - (5); DM ec, TR ff; 7-21 Jul, 5 Oct-17 Nov; mt. Acropteroxys gracilis (Newman) - (3); DM tm and ec; 28 May-8 Aug; mt. *Tritoma biguttata biguttata (Say) - (abundant); FH dp, GF qu, TR ff; 21 May-10 Oct; hp (in gills of yellowish- *Ischyrus quadripunctatus (Olivier) - (19); GF ff, TR ff orange Russula sp. [Russulales: Russulaceae; near R. and uf; 10 Apr-4 Sep; mt, uv. ochroleucoides Kauffm]), mt. Languria angustata (Beauvois) - (3); DM tm; 16 Jun-8 *Tritoma biguttata affinis Facordaire - (8); FH dp, TR Aug; mt. ff; 5 Aug-10 Oct; mt. Languria mozardi Latreille - (1); FM uf; 15 Apr; bf. This subspecies is reported by Evans (2014) as not occurring on the East Coast north of southern North Loberus impressus FeConte - (8); DM tm; 11 Apr-1 Carolina, however Downie & Arnett (1996) list records Aug; mt. from as far north as Pennsylvania. *Megalodacne fasciata (Fabricius) - (2); TR uf, FH dp; *Tritoma erythrocephala Facordaire - (2); LH dp, 11 Jun-30 Jul; hp (shelf fungus on stump), mt. TR ff; 7-21 Jun, 19 Sep-10 Oct; mt. NEW STATE RECORD. *Megalodacne heros (Say) - (4); GF uf; 24 Jun-6 Jul; hp (on fungus), uv. This beetle was expected in Virginia because it was documented previously in many states from Texas and * Micro sternus ulkei (Crotch) - (2); TR ff, GF sf; 10 Apr- Kansas eastward to New York and Florida (Downie & 21 Jun; mt. NEW STATE RECORD. Arnett, 1996). This species has a narrow range in North America *Tritoma humeralis Fabricius - (abundant); DM ff, FH extending from Illinois (Goodrich, 1994) and Tennessee dp, GF sf, TR ff; 21 May-10 Oct; pf, mt. eastward to Pennsylvania and North Carolina (Downie & Arnett, 1996). It appears to have a relict distribution, A Malaise trap set from 19 September-10 October being the only Nearctic representative of a widely 2017 at Fittle Hunting Creek contained 88 T. humeralis, distributed genus. It is not common where it occurs 34 T. b. biguttata, 7 T. b. affinis, and 1 T. erythrocephala. (Boyle, 1956). It has been documented on shelf fungi in the family Hymenochaetacea (Goodrich, 1994). *Tritoma mimetica (Crotch) - (4); GF ff and qu, TR ff; 23 May-30 Jun; mt. NEW STATE RECORD. *Toramus pulchellus (FeConte) - (8); AW uf, GF ff; 15 Apr, 23 Jun-6 Aug; bf, bl, uv. Virginia is well within the range of this species, which has been documented from Texas and Kansas *Triplax festiva Facordaire - (5); AW uf, FH dp, GF sf; eastward to Quebec and Florida (Downie & Arnett, Apr 10-30, 3 Jun-15 Oct; bf, If, mt. 1996). *Triplax flavicollis Facordaire - (abundant); DM ec, GF *Tritoma pulchra Say - (6); GF ff, TR ff; 1 May-26 Jul; qu, TR uf; 17 May-17 Nov; hp (under bark and on mt. fungus), mt. *Tritoma sanguinipennis (Say) - (10); LH dp, GF sf and A Malaise trap set in floodplain forest in Turkey Run qu, TR ff; 10 Apr -30 Jul; mt. Park from 22 October-17 November 2008 contained 63 T. flavicollis and 2 T. thoracica. *Tritoma unicolor Say - (4); GF sf; 19-30 Jun; mt. 26 BANISTERIA NO. 50, 2018 Tetratomidae (Polypore Fungus Beetles) *Eustrophus tomentosus Say - (4); GF riverside prairie and uf, TR uf; 15 Jan, 14 Apr; hp (under bark and under This family was split from the Melandryidae (false loose bark of dead standing Quercus coccinea Miinchh. darkling beetles) by Nikitsky (1998). Evans (2014) [Fagales: Fagaceae]). reported 17 species in the eastern United States and 12 species are reported for South Carolina (Ciegler, 2014b). * Hallomenus scapularis Melsheimer (Figs. 2-3) - (4); Nine species have been documented at GWMP. TR uf; 5 Sep-21 Oct; mt. NEW STATE RECORD. *Eustrophopsis bicolor (Fabricius) - (4); DM ec and ff, Pale form (perhaps subteneral) Hallomenus TR ff, GF ff; 19-28 Apr, 30 Jun-13 Jul, 11-27 Sep, 15 scapularis (Fig. 2) is very similar to descriptions of Dec; hp (under bark of dead standing Carya tomentosa H. debilis FeConte. These two species overlap in range [Fam.] Nutt) (Fagales: Juglandaceae), mt. Fig. 2. Hallomenus scapularis Melsheimer (pale form); Collection data: Virginia, Fairfax Co., Turkey Run Park, gulch, Malaise trap, 5 September-21 October 2009, B. Steury & D. Smith. Fength 4.0 mm. The slightly paler humeral angles Fig. 3. Hallomenus scapularis Melsheimer (typical form). can be seen even in pale form specimens. Same collection data as Fig. 2. STEURY: FUNGIVOROUS BEETLES 27 and body length. They are separated in keys (Downie & Brown, J. W. (ed.). 2008. The invertebrate fauna of Arnett, 1996; Ciegler, 2014b) based on dorsal coloration Plummers Island, Maryland. Contribution XXX to the rather than anatomical features. The two can reportedly Natural History of Plummers Island, Maryland. Bulletin be separated by the extent of the basal punctures on the of the Biological Society of Washington 15: 1-226. pronotum; “small depressions” in H. debilis and “large, deep impressions” in H. scapularis (Ciegler, 2014b). Ciegler, J. C. 2014a. South Carolina beetles: Erotylidae These four specimens were captured in the same Malaise (http://scbeetles.info). (Accessed 5 June 2018.) trap, but only one was of the pale form. Ciegler, J. C. 2014b. Tenebrionoidea of South Carolina. *Holostrophus bifasciatus (Say) - (16); DM ec, LH dp, Biota of South Carolina. Volume 8. Clemson University, GF sf, TR uf; 10 Apr-20 Jul; pf, mt. Clemson, S.C. 244 pp. *Penthe obliquata (Fabricius) - (5); GF sf, TR ff; 16 Downie, N. M., & R. H. Arnett, Jr. 1996. The Beetles of Jun-17 Aug; hp (under bark), If, mt. Northeastern North America. Volume II. Sandhill Crane Press, Gainesville, FL. 830 pp. *Penthe pimelia (Fabricius) - (7); LH dp, GF sf and uf, TR ff and uf; 10 Apr-24 Aug; hp, If, pf, mt. Evans, A. V. 2008. Beetles. Pp. 44-51 In A. V. Evans (ed.), The 2006 Potomac Gorge Bioblitz - overview and *Pisenus humeralis (Kirby) - (1); TR ff; 22 Oct-17 Nov; results of a 30-hour rapid biological survey. Banisteria mt. 32. *Synstrophus repandus (Horn) - (15); GF riverside Evans, A. V. 2014. Beetles of Eastern North America. prairie, TR uf and ff; 14 Apr, 24 Jun-21 Jul; bs, hp (under Princeton University Press, Princeton, NJ. 560 pp. bark), mt. Gandhi, K. J. K., & D. A. Herms. 2010. North American Two specimens are of the brown form, the remainder arthropods at risk due to widespread Fraxinus mortality are of the more typical black form. caused by the alien emerald ash borer. Biological Invasions 12: 1839-1846. *Tetratoma tessellata (Melsheimer) - (1); GF ff; 21 May-18 Jun; mt. Goodrich, M. A. 1994. Microsternus ulkei (Crotch) (Coleoptera: Erotylidae), a western extension of its range ACKNOWLEDGMENTS and first record from Illinois. Transactions of the Illinois State Academy of Science 87: 171-174. Appreciation is extended to our Bug Lab volunteers, M’Shae Alderman, Judy Buchino, Pat Findikoglu, Kukor, J. J., & M. M. Martin. 1987. Nutritional ecology Peggy Finn, Tom Hahn, Sarah Hill, Ann Kelly, Eileen of fungus-feeding arthropods. Pp. 719-814 In F. Slansky Miller, Susan Sprenke, and Jerry Taylor for their & J. G. Rodrigues (eds.), Nutritional Ecology of Insects, assiduous sorting of Malaise trap samples and to Sam Mites, Spiders, and Related Invertebrates. John Wiley & Droege and Erick Hernandez, Bee Inventory and Sons, New York. Monitoring Laboratory, Patuxent Wildlife Research Center, United States Geological Survey, for producing Lawrence, J. F. 1989. A catalog of the Coleoptera of all of the images except Figure 1 bottom. Steve Roble, America north of Mexico, family Derodontidae. United Virginia Department of Conservation and Recreation, States Department of Agriculture. Agriculture Division of Natural Heritage, and two anonymous Handbook Number 529-65. 14 pp. reviewers provided very helpful comments on the draft manuscript. Nikitsky, N. B. 1998. Generic classification of the beetle family Tetratomidae (Coleoptera, Tenebrionoidea) of LITERATURE CITED the world, with description of new taxa. Pensoft Series Faunistica 9, Sofia, Bulgaria. 80 pp. Boyle, W. W. 1956. A revision of the Erotylidae of America north of Mexico (Coleoptera). Bulletin of Robertson, J. A., A. Slipinski, M. Moulton, F. W. the American Museum of Natural History 110: 61- Shockley, A. Giorgi, N. P. Lord, D. D. McKenna, W. 172. Tomaszewska, J. Forrester, K. B. Miller, M. F. Whiting, 28 BANISTERIA NO. 50, 2018 & J. V. McHugh. 2015. Phylogeny and classification of beetles of the world (Coleoptera: Cucujoidea: Cucujoidea and the recognition of a new superfamily Endomychidae). Zootaxa 1999: 1-113. Coccinelloidea (Coleoptera: Cucujiformia). Systematic Entomology 40: 745-778. Steury, B. W. 2011. Additions to the vascular flora of the George Washington Memorial Parkway, Virginia, Schigel, D. S. 2008. Collecting and rearing fungivorous Maryland, and the District of Columbia. Banisteria 37: Coleoptera. Revue d’ecologie - la Terre et la Vie 63: 7- 3-20. 12. Steury, B. W., G. P. Fleming, & M. T. Strong. 2008. Schigel, D. S. 2012. Fungivory and host associations of An emendation of the vascular flora of Great Falls Park, Coleoptera: a bibliography and review of research Fairfax County, Virginia. Castanea 73: 123-149. approaches. Mycology 3: 258-272. Townes, H. 1962. Design for a Malaise trap. Shockley, F. W., K. W. Tomaszewska, & J. V. McHugh. Proceedings of the Entomological Society of 1999. An annotated checklist of the handsome fungus Washington 64: 253-262.

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