Number 53 BANISTERIA A JOURNAL DEVOTED TO THE NATURAL HISTORY OF VIRGINIA 5 She, Styeip frre baecis rus Ly ‘ (exe ado erecto ste ineluses. Xe ISSN 1066-0712 2019 BANISTERIA A JOURNAL DEVOTED TO THE NATURAL HISTORY OF VIRGINIA ISSN 1066-0712 Published by the Virginia Natural History Society The Virginia Natural History Society (VNHS) is a nonprofit organization dedicated to the dissemination of scientific information on all aspects of natural history in the Commonwealth of Virginia, including botany, zoology, ecology, archeology, anthropology, paleontology, geology, geography, and climatology. Membership in VNHS includes a subscription to Banisteria. Annual dues are $20.00 (per calendar year); library subscriptions to Banisteria are $40.00. Checks should be made payable to the Virginia Natural History Society; an online payment method is also available. Membership dues and inquiries should be directed to the Co-Treasurers (address, page 2); correspondence regarding Banisteria to the Editor. Banisteria 1s a peer-reviewed journal. The Editor will consider manuscripts on any aspect of natural history in Virginia or neighboring states if the information concerns a species native to Virginia or the topic is directly related to regional natural history (as defined above). Biographies and historical accounts of relevance to natural history in Virginia also are suitable for publication in Banisteria. For additional information regarding the VNHS, including other membership categories, annual meetings, field events, pdf copies of papers from past issues of Banisteria, and _ instructions for prospective authors, consult our website at: http://virginianaturalhistorysociety.com/ Editorial Staff: Banisteria Editor Todd Fredericksen Ferrum College 215 Ferrum Mountain Road Ferrum, Virginia 24088 Production Consultant Steven M. Roble Banisteria No. 52 and 53 were published on 15 January 2020. Cover: Magnolia virginiana Linnaeus (Sweetbay); original drawing by John Banister, sent to Bishop D. H. Compton in 1689. Figure 90 in folio in Sir Hans Sloane’s MS 4002 in the British Museum. Back cover: Anemone quinquefolia Linnaeus (Wood Anemone). Original drawing by John Banister; sent to Bishop D. H. Compton in 1689. Figure 72 in folio in Sir Hans Sloane’s MS 4002 in the British Museum. Photocopies courtesy of the late Joseph and Nesta Ewan. BANISTERIA A JOURNAL DEVOTED TO THE NATURAL HISTORY OF VIRGINIA Number 53, 2019 Contributed Papers Natural History of the Eastern Harvest Mouse in Southeastern Virginia IRGD ETE IY IROSEs MU tee de cdrtete ase lea german trncidiacers tiileatebyagutesrderdetarmicc defi pretmmdenntedenghiteansateony te of omntuectitanah beaten AP 3 Investigating Campus Features that Influence Bird-Window Collisions at Radford University, Virginia Karen E. Powers, Lauren A. Burroughs, Breann M. Mullen, Hannah C. Reed, and Zoe Q. Krajcirovic .............. 11 Using DNA Barcoding to Identify Carcasses from Bird-Window Collisions at Radford University Claudia Y. Paniagua-Ugarte, Karen E. Powers, and Robert R Sheehy.................000000ccccceccececcecseeceeeesesseeeeeennees De The Rove Beetles (Coleoptera: Staphylinidae) of the George Washington Memorial Parkway, with a Checklist of Regional Species R. Michael Brattain, Brent W. Steury, Alfred F. Newton, Margaret K. Thayer, and Jeffrey D. Holland.............. 27 Probable Cerulean Warbler x Northern Parula Hybrid in Rockbridge County, Virginia in April 2019 Richard. Rowe ands ieincda Wi RO Wee... i ecncnat cog sperteereiotaalee icon Rendbce sy eunedianneyad nore. Dg ragh hse recent eae: 72 First Records of the Neotropical Deer Ked Lipoptena mazamae Rondani (Diptera: Hippoboscidae) from Virginia Liberty Hightower, Nancy D. Moncrief, and Kaloyan Ivanov.........0..0.. 0.0 c ccc cece tne naes 78 Notes on the Parasitic Beaver Beetle, Platypsyllus castoris Ritsema, 1869 and Mouse Nest Beetle, Leptinus orientamericanus Peck, 1982 (Coleoptera: Leiodidae: Platypsyllinae) in Virginia Ralph Reker atid Atta uri ye BVA Sct ca sbyacclte eee idee boilaentte ster treeton ees d/da unneatloc onteetvnsiceee eee tel mmmngel tenes trae, 84 Miscellanea ] Se) 0161 ean Topp pom Aaa RA WOE Sede As 29 none ih NN REN oo oa Top ow FRA. WOE RE St deh = As 9 ines eh ARE AED ane Deed A ba Ae Oe 87 TUNE IEC CIM CTIT STO Po oo EAP ds LLL Ste oes BP CRESS SUE BGR Sons EER Adds LL Le Geren skeet Bc AALS IO RAE SUS Noah ae 89 SUICeMt WeniDErsilpINCEMtIVGy:?, cass Pernice set ay.teoeeesceot secnaseeatnn tek ee eouenccans ouster erst it ag yuishiivacsecoasdeeuonatreet rit 89 The Virginia Natural History Society — Articles of Incorporation and Bylaws ........0..0..cccccccccccceecccteeeeenees 90 Virginia Natural History Society Officers, 2019 President Nancy Moncrief Virginia Museum of Natural History Martinsville, Virginia 24112 nancy.moncrief@vmnh.virginia.gov (term expires December, 2020) Vice President Kal Ivanov Virginia Museum of Natural History Martinsville, Virginia 24112 kal.1vanov@vmnh. virginia. gov (term expires December, 2020) Co-Treasurers Nancy Moncrief and Kal Ivanov Virginia Museum of Natural History Martinsville, Virginia 24112 (terms expire December, 2022) Secretary and Webmaster Paul Marek Department of Entomology Virginia Tech Blacksburg, VA 24061 pmarek@vt.edu (term expires December, 2021) Councilors Karen Powers, Radford, VA (term expires December, 2021) Arthur Evans, Richmond, VA (term expires December, 2022) Curt Harden, Clemson, SC (term expires December, 2022) Banisteria, Editor Steven Roble (through issue #52) steve. roble@dcr. virginia. gov Todd Fredericksen (beginning with issue #53) tfredericksen@ferrum.edu Honorary Councilors Michael Kosztarab, Joseph Mitchell Banisteria, Number 53, pages 3—10 © 2019 Virginia Natural History Society Natural History of the Eastern Harvest Mouse in Southeastern Virginia Robert K. Rose Department of Biological Sciences Old Dominion University Norfolk, Virginia 23529-0266 brose@odu.edu ABSTRACT The Eastern Harvest Mouse, Reithrodontomys humulis, has been studied extensively in southeastern Virginia since 1979, using a combination of live and pitfall trapping methods. This smallest rodent of eastern North America also is one of most versatile, occupying a range of habitats in southeastern Virginia from old fields in different stages of succession, brushy edges, and forests of different types. As with other species of Reithrodontomys, R. humulis often is associated with the Hispid Cotton Rat, Sigmodon hispidus, with both reaching modest densities in old fields. Two capture-mark-release studies of small mammal communities in southern Chesapeake lasting eight and nine years revealed that the Eastern Harvest Mouse was third in total abundance, behind Hispid Cotton Rat and Meadow Vole, Microtus pennsylvanicus, as old fields transitioned into forests. Multiple field studies using pitfall traps in a range of habitats in southeastern Virginia also indicated that harvest mice often arrive early in succession and stay later than other rodents. Keywords: Coastal Plain, Eastern Harvest Mouse, habitat selection, Reithrodontomys humulis, small mammals. INTRODUCTION The Eastern Harvest Mouse, Reithrodontomys humulis, is a cricetid rodent with a distribution mostly in the southeastern U.S. (Stalling, 1997). With adults averaging about 8 g, this is the smallest rodent in the eastern U.S. Its small size alone distinguishes it from the 16-25 g White-Footed Mouse, Peromyscus leucopus, the native rodent with which adults are most comparable in coloration and body form; both have brownish backs, white or nearly white underbellies, and long tails. The other similar small mouse with which R. humulis might be compared 1s the House Mouse, Mus musculus, which has large and naked ears, a nearly hairless unicolored tail, and a gray or orangish underbelly. Their behaviors differ too; when placed in a bucket after removal from a live trap, an Eastern Harvest Mouse is likely to remain calm and groom itself or eat seeds, whereas a House Mouse is frenetic, running and leaping in its attempts to escape. Much of the information in this report comes from the work of Old Dominion University graduate students conducting field research projects as part of their thesis research for the Master of Science degree. Jean Ferguson Stankavich, who conducted capture-mark-release (CMR) studies of small mammals in the northwest section of the Great Dismal Swamp National Wildlife Refuge, found Eastern Harvest Mice to be numerically dominant in two CMR grids. Sarah Crawford added an analysis of vegetation composition and structure to her study of small mammal communities with harvest mice. Michelle Cawthorn Chandler used a specially built trap, with a 2.1 by 2.1 cm opening, to exclude larger small mammals in an effort to study the smallest members of the small mammal community in an old field. These studies were conducted in habitats in early stages of succession, where densities of small mammals tend to be highest. Additional information about distribution and relative abundance comes from field studies using pitfall traps on dozens of 0.25 ha grids that enable comparisons 4 BANISTERIA of relative densities among habitat types and from two long-term CMR studies of rodent communities. GENERAL CHARACTERISTICS The Eastern Harvest Mouse is grayish brown with a darker mid-dorsal stripe on the back, with lighter and sometimes rusty sides, and whitish feet (Fig. 1). The tail is about the same length as the head-body length; in a series of harvest mice from Isle of Wight County measured by the author, the tail was 47.8% of total length for 32 males and 47.6% of total length for 30 females. The underside of the tail is whitish, as is the belly. The eye is large and dark, suggesting nocturnal behavior. The vibrissae (whiskers) are numerous, long, and pale at the tips. Another feature that distinguishes the Eastern Harvest Mouse from other long-tailed rodents in southeastern Virginia is the groove in the upper incisor, which can be seen with the naked eye. The anterior face of the incisor is folded, giving it a corrugated appearance. The function of this feature is unknown, but the fold probably strengthens the tooth, thus reducing the likelihood that the tooth will break when opening hard- coated seeds. The sexes are similar in size (Dunaway, 1968), but when weighed with a 10-g Pesola scale (with 0.2 g calibrations) non-pregnant females from Chesapeake, Virginia (x = 8.20 + 0.3 SE g, n = 35) weighed significantly (p < 0.05) more than male harvest mice (x = 7.04 + 0.1 SE g, n = 42) (Cawthorn & Rose, 1989). The weight of males was relatively constant throughout the year but weights of females peaked in autumn, suggesting that as the season of greatest reproduction. Fig. 1. An adult eastern harvest mouse, Reithrodontomys humulis. Photo credit to West Virginia University Wildlife and Fisheries Science study guide (Edwards). NO. 53, 2019 DISTRIBUTION The Eastern Harvest Mouse has a_ mostly southeastern distribution in the US, extending eastward from eastern Oklahoma and Texas to states lying south of the Ohio River, but also including southern Ohio, western Maryland, all of Virginia, and points southward. It may be absent from the southern tip of Florida. Some populations in Texas, Oklahoma, Arkansas, and Louisiana overlap in distribution with those of R. fulvescens, a larger species that has been studied extensively in the Texas coastal prairies by Cameron (1977). Information on the distribution of R. humulis is accumulating as more community studies are being published, so the map of Stalling (1997), already an improvement of Hall (1981), continues to be revised. For example, before 1988, R. humulis was known from only three counties in Oklahoma and was considered a rare mammal, but by 2011, its presence had been recorded in six more counties (Braun et al., 2011). Three subspecies are recognized. Howell (1940) described R. humulis virginianus based on specimens from Amelia County, located in central Virginia just one county southwest of Richmond. This subspecies, present in the eastern half of Virginia, is paler and more grayish, with a blackish-brown mid-dorsal stripe, and with white feet compared to R. h. humulis, the other subspecies east of the Mississippi River. R. h. merriami is present in the four western states. As presently understood, the northern distribution of coastal populations of R. humulis is in southeastern Virginia. Field studies of the Eastern Shore of Virginia by Rose and colleagues (e.g., Rose & March, 2013) have failed to record any R. humulis in either Northampton or Accomack counties, so its movement northward likely is blocked by _ the Chesapeake Bay. Pagels & Moncrief (2015) also consider R. humulis to be absent on the Eastern Shore. FORM AND FUNCTION Its small size and long tail suggest that this rodent can climb into even herbaceous vegetation, perhaps to glean seeds or capture insects. Relatively little is known about its diet, except that it eats some seeds. But R. humulis is not considered to be truly arboreal because its softball-sized grassy nests are placed in low herbaceous vegetation or on the ground rather than in tree holes, as truly arboreal rodents usually do. The monthly mean weights of males from southeastern Virginia were relatively stable (Chandler, 1984). In Tennessee, unbred lab-reared adult harvest ROSE: EASTERN HARVEST MOUSE 5 mice of both sexes had identical weights, 8.2 g (Dunaway, 1968) and Kaye (1961) reported that lab- reared 50-day-old adult males and females weighed the same. By contrast, field-caught females were heavier than males in all but 3 of 21 months in Tennessee (Dunaway, 1968), suggesting that pregnancy accounts for most differences in weights of the sexes. As in southeastern Virginia, the weights of males were fairly constant throughout the year (Dunaway, 1968). In brief, adults are similar in size, about 8 g, and of equal body length. Small body size means that, on a per gram basis, R. humulis has a higher metabolic rate, and thus relatively higher energy requirements, than larger mammals, a relationship recognized decades ago by Kleiber (1961). Furthermore, below the temperature zone of least energy cost, termed thermoneutrality, the energetic costs increase disproportionately. For example, the resting metabolism of the Eastern Harvest Mouse at 23° C is 4.35 ml of oxygen per gram of body weight per hour, but at 7° C the metabolic rate more than doubles, to 9.62 ml of oxygen per gram per hour; the comparable values for the twice-as-large White-footed Mouse are 3.04 and 5.68 (Dunaway, 1968). Also, because of its small size, it can neither reduce heat loss via long and dense insulative fur nor accumulate large fat reserves, two ways larger mammals can conserve or produce heat during periods of cold temperatures. These factors likely restrict the distribution of Reithrodontomys, a genus with tropical origins, to sub-tropical and temperate climate zones in the US. The numbers of red blood cells per unit volume were similar to those of larger rodents examined by Dunaway (1968). However, harvest mice had much _ higher concentrations of hemoglobin (g/ml) in the erythrocytes than in larger rodents, likely an adaptation to deliver sufficient oxygen to cells of a small mammal with high metabolic rate. REPRODUCTION The breeding season for R. humulis likely varies by geographical location, starting earlier in spring in southern than in northern populations. In southeastern Virginia, breeding peaks were observed in spring and autumn, with a lull in summer (Cawthorn & Rose, 1989). The higher body weights of females plus the many gray- backed juveniles indicate greater levels of reproduction in autumn than in spring. Studies 1n the laboratory indicate that females in late pregnancy become intolerant of males and that males take no role in parenting (Kaye, 1961), the pattern seen in most mammals. Near the end of the 21-day gestation period, the female builds a birthing nest of dried grasses, in which the young are reared for about three weeks. Litter size averaged 2.2 for nine lab females in Florida (Layne, 1959) but was 3.2 for nine lab females from North Carolina (Kaye, 1961). Later, Dunaway (1962) reported finding three litters of three and three litters of four born in live traps in Tennessee; he also took a 17-g female into the lab where two days later it gave birth to eight young, the weights of which totaled nearly 8 g. Taken together, the litter size is about three; these are weaned near the end of the third week of life, at weights of about 5 g, the lightest animals trapped in most studies. On 16 December 2018, I recorded a 12-g pregnant female with partially open pubic symphysis and enlarged nipples, indicating that this female produced a late litter in southeastern Virginia. In my experience, females heavier than 10 g are pregnant. Few details are known about reproduction in male eastern harvest mice, in part because indicators of reproduction are fewer than in females. During the breeding season the enlarged testes are descended into the scrotum, and such males are judged to be reproductive. In the winter non-breeding season, the testes of many mammals, including harvest mice, decrease dramatically in size, often losing 95% of their weight, and such males are non-reproductive. Cawthorn & Rose (1989) observed scrotal males in every month of the year, with lowest rates (10%) in winter. In the nearby Great Dismal Swamp, Stankavich (1984) also found some scrotal males in winter (24%), suggesting the possibility of occasional year-round breeding in southeastern Virginia. Coastal Virginia averages 10 cm of snow, 10—20 nights below 0° C, and short periods of frozen soil. By contrast, no scrotal males were observed in December, January, and February in Tennessee (Dunaway, 1968). These observations suggest that populations in southern states (or coastal locations in Virginia) might have year-round reproduction, although it was not observed by Layne (1959) in Florida. In South Carolina, highest numbers of captures were recorded in January, indicating that breeding levels were greatest in late autumn (Briese & Smith, 1974). ECOLOGY Much new information about R. humilis in southeastern Virginia has been published in recent decades by using CMR methods on small square or rectangular (row by column) grids with live traps placed at the coordinates. At monthly or twice-monthly intervals, the traps are baited and ‘run’ for three consecutive days. Each captured animal is given a unique number, usually with an ear tag, weighed, and its sex and reproductive condition are recorded. The animal is then released at the point of capture. The goal is to trap 6 BANISTERIA such tagged animals in successive months and record the events of their lives: features such as their changes in body mass, levels of reproduction, rates of body growth and survival, area of use on the grid, among others. Also, the vegetation of the grid often is studied, both for its list of plant species but also for details of plant contacts at different heights in an effort to learn whether the vertical structure of the plant community is more or less important than the presence of certain plant species. For example, when grasses dominate the plant community, vertical structure is dense with stems and leaves below about 0.5 m. Later in biological succession, when shrubs and trees are common, the density of vegetation near the ground surface is much less, but vertical elements are more common, increasing vegetation complexity in a different way. The first CMR study in southeastern Virginia was conducted under a 40-m wide powerline in the northwest section of the Great Dismal Swamp National Wildlife Refuge (Stankavich, 1984). Two study grids of Fitch live traps (Rose, 1994) were established in habitat dominated by plants typical of early successional stages in a swamp: grasses and forbs, and in wetter places, sedges, rushes, and spikerushes. Some deciduous trees and shrubs were present too, especially in the slightly higher places where winter flooding did not kill them. Harvest mice were the most common small mammal in this habitat, comprising 71 of 155 total individuals (Rose & Stankavich, 2008). In an 18-month CMR study, conducted in Suffolk just west of the Great Dismal Swamp National Wildlife Refuge, seven harvest mice were tagged, along with 47 Synaptomys cooperi (Southern Bog Lemming) and 110 Microtus (Pitymys) pinetorum; no other rodents were captured in this community where minor species dominated (Rose & Ford, 2012). Michelle Cawthorn (Chandler, 1984) conducted CMR trapping of small mammals every other week for a year on two grids in an old field in the Bowers Hill region of Chesapeake. The tiny specially built traps excluded adults of the common small mammals and thus she caught mostly Eastern Harvest Mice and House Mice, 51% and 39%, respectively, of 703 total captures (Cawthorn & Rose, 1989). Highest densities for R. humulis were achieved on both grids in autumn and winter, with 44 harvest mice per hectare; the average density was 21.9 and 21.8/ha on both grids across the study. The adult mortality rate of 6 percent per month was constant for the year-long study. Home range, the area of greatest use, was similar for both sexes, at about 1000 m?. Lifespans, based on three or more captures, averaged about 10 weeks for both sexes, which if added to the 20-30 days for newborns to become trappable, equates to mean lifespans of about 100 days (Cawthorn & Rose, 1989), similar to those (90-120 days) in NO. 53, 2019 Tennessee (Dunaway, 1968). Cawthorn/Chandler, (1984) recorded 29 plant species on Grid 1 and 27 species on Grid 2, 18 of which were present on both grids; asters dominated on Grid 1 but honeysuckles (Lonicera) on Grid 2. But height of vegetation was more important than species composition, a conclusion also reached by Crawford (2013), who used assessments of plant composition and measurements of structure while trying to understand the strong association of harvest mice with the Hispid Cotton Rat (Sigmodon hispidus), adults of which are mostly 80— 120 g in southeastern Virginia. Numerous studies report that S. hispidus and the local Reithrodontomys species often occur together, regardless of the region. Both are tropical genera with populations in the US at the northern limits of distribution (e.g., Braun et al. [2011]; Brady & Slade [2001] for R. megalotis in eastern Kansas; Rose et al. [2018] for R. humulis in southeastern Virginia). Both genera reach highest densities in old field and other early successional habitats, but the reasons for their frequent coexistence remain unclear, whether by being active at different times of day, by mutual avoidance at the microhabitat level, or by differential use of resources (Crawford, 2013). Using live-trapping records, Crawford (2013) found no evidence that either harvest mice or cotton rats avoided the other on either of two 1-ha grids, each trapped monthly for multiple years. A negative association between captures at each station was recorded for only one month over that period. Both species tended to occupy areas with few or no trees, and harvest mice were more likely than cotton rats to be present if the open sites were wet. Both species tended to use areas with dense vegetation near the ground surface, regardless of plant species composition. Crawford speculated that differential use of resources (harvest mice are primarily seed-eaters whereas cotton rats eat stems and leaves, mostly of monocots) and the broader habitat tolerances of harvest mice as the most likely reasons for the coexistence of these two species in southeastern Virginia. In evaluating the changes in composition of the community of small mammals on the same two grids analyzed by Crawford (2013), Rose et al. (2018) found that harvest mice and cotton rats were early colonizers in grassy old fields in the third year after a farm field was abandoned and both species persisted while other community members came and went. Across eight years of study on one site and nine years on the other, R. humulis was third in total abundance on both grids, with cotton rats being most numerous on one grid and Meadow Voles (Microtus pennsylvanicus) numerically dominant on the other. Thus, although many investigators would consider harvest mice to be a minor ROSE: EASTERN HARVEST MOUSE 7 species in the community of small mammals, in southeastern Virginia they are early arrivals, third in abundance during succession, and they are still present at the point when the forest small mammals, such as White-footed Mice and Golden Mice (Ochrotomys nuttalli), arrive and become the dominant rodents. In field studies using pitfall traps on 0.25 ha grids, the results were similar. For example, R. humulis was present on 13 of 14 grids in Isle of Wight County, more than any other species, and was second in abundance to Least Shrew (Cryptotis parva) (Rose, 2005). Similar results were found in 19 pitfall grids in Virginia Beach, Chesapeake, and Suffolk (Rose, 2016). In his pitfall-trapping study of small mammals in and near the Great Dismal Swamp National Wildlife Refuge, Everton (1985) found R. humulis on 10 of 21 one-quarter hectare grids, and fourth in overall abundance behind two shrews (Southeastern Shrew, Sorex longirostris, and Least Shrew) and Southern Bog Lemmings (Synaptomys cooperi). In the analysis of vegetation structure, Everton found that R. humulis was associated with high values for stem densities from ground level to 40 cm and for average height of herbaceous vegetation, indicating a strong preference for dense cover of plants, mostly grasses, near the surface. In a summary of studies of small mammals conducted across the range of habitats in the Great Dismal Swamp, using all trapping methods (live, pitfall, and break-back traps [used in the late 19" century]), R. humulis was third in total abundance, behind Short-tailed Shrews (Blarina spp.) and White-Footed Mouse (Rose et al., 1990, Table 4). Thus, in southeastern Virginia at least, R. humulis is the most versatile rodent in the small mammal community. For example, one was caught on a tall sand dune at Little Creek Amphibious Base in Norfolk, along with House Mice and White-footed Mice (Rose & Sweitzer, 2013). More commonly, R. humulis arrives early in old fields dominated by grasses and forbs, sometimes sharing early arrival status with house mice. Soon other species, such herbivores as cotton rats, meadow voles, and rice rats, arrive and some of these become dominant species for months or years. But when the herbaceous vegetation thins and eventually is shaded out by shrubs, saplings, and trees, the herbivorous rodents disappear, often quickly. Based on studies of two old fields going through succession, cotton rats and harvest mice often were still present before forest rodents come to dominance. Eastern Harvest Mice are much less common in the forests of southeastern Virginia than in earlier stages of succession, but often they are present in small numbers (e.g., Everton, 1985). Others also have found R. humulis in forests, such as in wetland forests in Tombigbee National Forest in Mississippi (Edwards & Jones, 2014), and rarely in pine forests (Dolan & Rose 2007). In their pitfall trapping study in the upper coastal plain of Virginia, Bellows et al. (2001) found R. humulis to be more abundant in old field habitats than in other macrohabitats; harvest mice were present in oak-hickory forest and young pine forests, but not in older pine or oak-pine forests. In a four-year study in the North Carolina coastal plain, R. humulis had good recruitment and survival in all five treatments that provided varying amounts of structure (woody debris, pine seedlings, switchgrass), and by year four it outnumbered the other three colonizing species (Homyack et al., 2014). In brief, numerous studies reveal R. humulis to be versatile by occupying a range of habitat types. BEHAVIOR Harvest mice are primarily nocturnal, and thus are active during the coldest part of the day, enabling them to benefit from the heat generated during foraging and other activities. At thermoneutrality (22° C), R. humulis shows an innate increase in metabolic rate at the approach of darkness, as if foretelling the beginning of nocturnal behavior (Baker, 1974). Baker, who measured CO production rather than oxygen consumption, also recorded a doubling of metabolic rate when harvest mice were housed at 9° C. Nocturnal behavior means that owls are their main avian predators, as recorded by Klippel & Parmalee (1982) in their study of pellets from a wintering Long- eared Owl (Asio otus) in the Nashville Basin of Tennessee. R. humulis was second in abundance (n= 78) to Prairie Vole (Microtus ochrogaster, n = 129) among 71 complete pellets. In a study conducted near Williamsburg, Virginia, Rosenburg (1986), who followed tagged Barn Owls (7yto alba) via radio- tracking, found small numbers of Eastern Harvest Mice in their pellets in most seasons. The Meadow Vole, also common in old fields in early succession, was the main food of these owls. The observation of multiple captures in live traps suggests some degree of sociality in harvest mice. In southeastern Virginia, 6.4% of total captures were as multiple captures, with more male-male pairs and fewer female-female pairs than expected (Cawthorn & Rose, 1989). Others have reported huddling, especially during winter months. For example, Dunaway (1968) reported that only 3 of 18 eastern harvest mice were alone in the nest cans of traps on a late January day; the others were in groups of 6, 4, 3, and 2. I have observed similar social groupings in R. megalotis in eastern Kansas, instances in which up to 11 adults shared grassy nests 1n gallon-sized nest chambers. Similar social groupings were observed 8 BANISTERIA in R. fulvescens in the Texas coastal plain (Spencer et al., 1982). Formation of social groups is especially important for tiny mammals, enabling them to share the costs of staying warm together in their well-insulated grass nests. In his analysis of spacing behavior among individuals of R. humulis, Dunaway (1968) found little evidence of territoriality: territorial individuals are anti- social. One consequence of social groupings is_ the potential for the ‘sharing’ of ectoparasites. Clark & Durden (2002) found 10% prevalence both of fleas (Polygenis gwyni) and of ticks (Amblyoma maculatum) in Eastern Harvest Mice in Mississippi. In southeastern Virginia, of nine small mammal species evaluated for ticks over a period of years, harvest mice had the lowest proportion of infestation; 18.3% had ticks, mostly on the ears (17 of 93; H. Gaff, pers. comm.). By contrast, another benefit of social groupings is allogrooming, 1.e., the removal of ectoparasites by other members of the group. There is no direct evidence of allogrooming in R. humulis, but the low percentage of ticks on harvest mice is consistent with this hypothesis. The killing of young by siblings or mother seems to be a common behavior, at least in captivity; sometimes this unexplained behavior is followed by cannibalism (Dunaway, 1962; Kaye, 1961). GENETICS Information on the chromosomes of R. and its congers is mostly derived from studies conducted nearly 30 years ago. Carleton & Myers (1979) reported that R. humulis had a diploid number of 2n = 51 for two females (no males were assessed); the chromosomes were mostly small-to-medium acrocentrics plus five pairs of larger and bi-armed chromosomes. The unpaired element was a small metacentric chromosome. The 2n = 51 was confirmed by Robbins & Baker (1980), and although they determined the FN to be 78, they could not determine the origin of the unpaired element. Much remains to be learned about the genetics of R. humulis and others in this genus. CONSERVATION STATUS The 2016 International Union for the Conservation of Nature Red List of Threatened Species lists R. humulis as “Least Concern,’ and with stable populations. However, a map on the same website indicates that Oklahoma considers the species to be “critically imperiled,” but this statement conflicts with Braun et al. (2011), which adds six counties to their known locations in the state. The map also lists the species as “possibly extirpated” in West Virginia and “not ranked” or “under NO. 53, 2019 review in Mississippi, South Carolina, and Florida. The Ohio Department of Natural Resources website states that R. humulis is a “Threatened” species in their state, despite Harder et al. (2014). The other states, including Virginia, assess their populations to be “secure” or “apparently secure.” REMARKS The name Reithrodontomys humulis was given in 1841 by John James Audubon and his son-in-law, John Bachman, based on specimens collected near Charleston, South Carolina. Early in the next decade, these same authors, far better remembered for their studies of and naming of many North American birds, published Quadrupeds of North America, the first comprehensive book on New World mammals. They chose the genus name, Reithrodontomys, derived from three Greek words (Lowery, 1974), because of the grooved incisor: reithron (groove), odous (tooth), and mys (mouse). The specific name humulis may be a misspelling of humilis, which means “little harvest mouse.” In their Quadrupeds book, the authors used the latter spelling. The tiny mouse of western Europe and the British Isles is also called “harvest mouse,” but it is ina different genus, Micromys, literally “tiny mouse.” ACKNOWLEDGMENTS I gratefully acknowledge the assistance of many former students while conducting small mammal studies in southeastern Virginia, especially Michelle Cawthorn (Chandler) and Sarah Crawford, both of whom focused their MS theses on understanding the ecology of Reithrodontomys humulis. Thanks also to other students who participated in field studies, including James Dolan, Jana Eggleston, Roger Everton, Jean Ferguson (Stankavich), Linda Ford, Heather Green (Salamone), Jay Kiser, Robyn Nadolny, Thomas Padgett, and John Rose. LITERATURE CITED Baker, C. E. 1974. Measurement of small mammal metabolism by infrared gas analysis. Journal of Mammalogy 55: 664-670. Bellows, A. S., J. F. Pagels, & J. C. Mitchell. 2001. Macrohabitat and microhabitat affinities of small mammal in a fragmented landscape on the upper coastal plain of Virginia. American Midland Naturalist 146: 345-360. Brady, M. J., & N. A. Slade. 2001. Diversity of a ROSE: EASTERN HARVEST MOUSE 9 grassland rodent community at varying temporal scales: the role of ecologically dominant species. Journal of Mammalogy 82: 974-983. Braun, J. K., L. J. Vitt, J. P. Caldwell, M. A. Mares, & M. A. Revelez. 2011. Mammals from LeFlore County, Oklahoma. Southwestern Naturalist 56: 410-417. Briese, L. A., & M. H. Smith. 1974. Seasonal abundance and movement of nine species of small mammals. Journal of Mammalogy 55: 615-629. Cameron, G. N. 1977. Experimental species removal: demographic responses by Sigmodon hispidus and Reithrodontomys fulvescens. Journal of Mammalogy 58: 488-506. Carleton, M. D., & P. Myers. 1979. Karyotypes of some harvest mice, genus Reithrodontomys. Journal of Mammalogy 60: 307-313. Cawthorm, M., & R. K. Rose. 1989. The population ecology of the eastern harvest mouse (Reithrodontomys humulis) in southeastern Virginia. American Midland Naturalist 122: 1-10. Chandler, M. Cawthorn. 1984. Life history aspects of Reithrodontomys humulis in southeastern Virginia. M.S. thesis, Old Dominion University, Norfolk, VA. 56 pp. Clark, K. L., & L. A. Durden. 2002. Parasitic arthropods of small mammals in Mississippi. Journal of Mammalogy 83: 1039-1048. Crawford, S. A. 2013. Friends or foes: interpreting the relationship between two syntopic small mammals in southeastern Virginia, the hispid cotton rat (Sigmodon hispidus) and eastern harvest mouse (Reithrodontomys humulis). M.S. thesis, Old Dominion University, Norfolk, VA. 89 pp. Dolan, J. D., & R. K. Rose. 2007. Depauperate small mammal communities in managed pine plantations in eastern Virginia. Virginia Journal of Science 58: 147— 163. Dunaway, P. B. 1962. Litter-size record for eastern harvest mouse. Journal of Mammalogy 43: 428-429. Dunaway, P. B. 1968. Life history and populational aspects of the eastern harvest mouse. American Midland Naturalist 79: 48-67. Edwards, K. E., & J. C. Jones. 2014. Trapping efficiency and associated mortality of incidentally captured small mammals during herpetofaunal surveys of temporary wetlands. Wildlife Society Bulletin 38: 530-535. Everton, R. K. 1985. The relationship between habitat structure and small mammal communities in southeastern Virginia and northeastern North Carolina. M.S. thesis, Old Dominion University, Norfolk, VA. 76 pp. Hall, E. R. 1981. The Mammals of North America. 2nd Edition. John Wiley & Sons, New York. 1,181 pp. Harder, J. D., J. K. Kotheimer, & I. M. Hamilton. 2014. A regional study of diversity and abundance of small mammals in Ohio. Northeastern Naturalist 21: 210- 233. Homyack, J. A., K. E. Lucia-Simmons, D. A. Miller, & M. Kalcounis-Rueppell. 2014. Rodent population and community responses to forest-based biofuel production. Journal of Wildlife Management 78: 1425-1435. Howell, A. H. 1940. A new race of the harvest mouse (Reithrodontomys) from Virginia. Journal of Mammalogy 21: 346. Kaye, S. V. 1961. Laboratory life history of the eastern harvest mouse. American Midland Naturalist 66: 439— 451. Kleiber, M. 1961. The Fire of Life: An Introduction to Animal Energetics. John Wiley & Sons, Inc., New York. A5A4 pp. Klippel, W. E., & P. W. Parmalee. 1982. Prey of a wintering long-eared owl in the Nashville Basin, Tennessee. Journal of Field Ornithology 53: 418-420. Layne, J. N. 1959. Growth and development of the eastern harvest mouse, Reithrodontomys humulis. Bulletin of the Florida State Museum 4: 61-82. Lowery, G. H., Jr. 1974. The Mammals of Louisiana and its Adjacent Waters. Louisiana University Press, Baton Rouge, LA. 565 pp. Pagels, J. F., & N. D. Moncrief. 2015. Virginia’s land mammals: past and present, with some thoughts about their possible future. Virginia Journal of Science 66: 171-222. 10 BANISTERIA Robbins, L. W., & R. J. Baker. 1980. G- and C-band studies on the primitive karyotype for Reithrodontomys. Journal of Mammalogy 61: 708-714. Rose, R. K. 1994. Instructions for building two live traps for small mammals. Virginia Journal of Science 45: 151— 157. Rose, R. K. 2005. The small mammals of Isle of Wight County, Virginia, as revealed by pitfall trapping. Virginia Journal of Science 56: 83-92. Rose, R. K. 2016. The small mammals of southeastern Virginia as revealed by pitfall trapping. Banisteria 47: 9-13. Rose, R. K., & L. J. Ford. 2012. Minor species as the dominant rodents in an oldfield. American Midland Naturalist 168: 1-8. Rose, R. K., & J. A. March. 2013. The population dynamics of two rodents in two coastal tidal marshes. Virginia Journal of Science 64: 17-26. Rose, R. K., & J. F. Stankavich. 2008. Low-density rodent communities in eastern Virginia. Virginia Journal of Science 59: 169-184. Rose, R. K., & J. L. Sweitzer. 2013. The small mammals of two dune communities in southeastern Virginia. NO. 53, 2019 Virginia Journal of Science 64: 151-157. Rose, R. K., R. K. Everton, J. F. Stankavich, & J. W. Walker. 1990. Small mammals in the Great Dismal Swamp of Virginia and North Carolina. Brimleyana 16: 87-101. Rose, R. K., R. M. Nadolny, J. Kiser, S. E. Rice, H. Green Salamone, J. Eggleston, & H. D. Gaff. 2018. Compositional changes in two small mammal communities during succession in southeastern Virginia. Virginia Journal of Science 69: 12 pp. Rosenburg, C. P. 1986. Barn owl habitat and prey use in agricultural eastern Virginia. M.S. thesis, College of William and Mary, Williamsburg, VA. 104 pp. Spencer, S. R., G. N. Cameron, & W. B. Kincaid. 1982. Multiple captures of the fulvous harvest mouse, Reithrodontomys fulvescens. American Midland Naturalist 107: 384-385. Stalling, D. T. 1997. Reithrodontomys humulis. Mammalian Species No. 565: 1-6. Stankavich, J. F. 1984. Demographic analysis and microhabitat relationships of a small mammal community in clearings of the Great Dismal Swamp. M.S. thesis, Old Dominion University, Norfolk, VA. 118 pp. Banisteria, Number 53, pages 11-21 © 2019 Virginia Natural History Society Investigating Campus Features that Influence Bird-window Collisions at Radford University, Virginia Karen E. Powers!', Lauren A. Burroughs, Breann M. Mullen, Hannah C. Reed, and Zoe Q. Krajcirovic Biology Department Radford University Radford, Virginia 24142 ‘Corresponding author: kpowers4@radford.edu ABSTRACT Window collisions pose a serious risk to birds, second only to domestic/feral cats. We sought to quantify the impacts of this threat at Radford University, a campus situated within a rural landscape and along a major migratory route (New River). We searched for evidence of bird-window collisions (BWCs) at 15 buildings in 2018 and 2019. In nearly 1,000 hours of surveys we discovered 51 birds (23 species) thought to result from BWCs. Increased window area tracked with a greater number of mortalities/building. Building height and surrounding vegetation metrics were not significantly related to BWCs. Species’ residency status did not significantly influence mortality events. Compared to BWC surveys nationwide, our number of mortalities was low, especially relative to our substantial surveying effort. Although this finding might suggest that Radford University buildings are not a significant source of mortality for birds, we recognize that (1) a priori surveying biases likely underestimated actual mortalities, and that (2) Radford University’s architectural changes in the last several years are increasing the likelihood of BWCs in the future. We suggest that Radford University explore window decals on current windows and incorporate “bird-friendly” glass on aspects that comprise large proportions of glass. Both of these steps contribute to Radford University’s goal of increasing the number of LEED-certified buildings on campus. Keywords: avian migration, avian mortality, building height, deterrents, time-of-day, vegetation, window area. INTRODUCTION Bird-window collisions (BWCs) are a substantial anthropogenic source of bird mortality, accounting for an estimated 365—988 million bird deaths annually. BWCs are the second largest cause of bird deaths, behind domestic and feral cats (Felis cattus, Loss et al., 2014: Kahle et al., 2016). Collisions typically are not limited to a particular avian taxon, and they can negatively affect common birds as well as species of conservation concern (Loss et al., 2014; Hager et al., 2017). Previous BWC studies have covered the gamut of building scenarios, from high-rises in a metropolis (Chicago: Briscoe & Dampier, 2019; Manhattan: Gelb & Delacretaz, 2009) to myriad college campuses (Hager et al., 2017). Multiple studies have investigated landscape and geographic metrics, as well as species-specific natural history features that may significantly affect the likelihood of BWCs. Features of the buildings, such as total window area and building height have been analyzed in several studies. While Bayne et al. (2012) found that collision rates were higher in rural areas where building density was lowest (Alberta, Canada), multiple studies found the opposite trend — higher building densities resulting in higher collision rates (Loss et al., 2014; Schneider et al., 2018). Hager et al. (2017) explained this difference along the spectrum of land development: building height and window area had a proportionally larger influence on BWCs in rural areas than in urban areas. This difference was most apparent during peak migration times, as non-resident birds were more likely selecting rural landscapes as suitable routes, and low-density buildings had a proportionally greater number of BWCs than buildings of the same size (height, window area) in an urban landscape. Artificial light also may increase BWCs, as nocturnal migrants may be 12 BANISTERIA confused by and attracted to them in flight (Hager et al., 2017). Features immediately surrounding buildings also have been studied as potential predictors of window collisions. Because surfaces like windows may reflect images of nearby vegetation, birds are likely confused, seeing it instead as a perching site or other suitable habitat (Hager et al., 2017). In many studies, vegetation effects were significant, but never primary metrics that influenced BWCs. Qualitative measures of vegetation have been implemented, but categories varied by study. The presence/absence (Chin, 2016) of vegetation was one method, while others loosely categorized density, using “some” vs. “extensive” vegetation (Gelb & Delacretaz, 2009) or “vegetated” vs. “less vegetated” (Gelb & Delacretaz, 2006). Quantitative vegetation metrics have covered the gamut of methods, but many used broad-scale vegetation metrics that relied on existing GIS-based layers. For example, Hager et al. (2017) utilized percent “vegetation” within 50 m of a building, while Schneider et al. (2018) examined percent landcover class within the same radius (e.g., lawn, trees, ornamental vegetation). Quantitative vegetation measures in situ were utilized less often, and were not always collected by the authors (e.g., Kummer et al. [2016a] asked citizen scientists to report the average height of vegetation around their residence). It is clear that no consistent method to measure vegetation exists for BWC studies. In these referenced studies, vegetation seems to have no more than a secondary effect on BWCs (typically less influential than window area or generalized building structure). Although avian and mammalian studies not focused on BWCs have successfully utilized metrics such as total vegetation volume and the Levins diversity index to quantify vegetation in situ (Francl & Schnell, 2002; Leighton et al., 2009), to our knowledge, no BWC studies have utilized these quantitative on-the-ground metrics to encapsulate vegetation effects. However, there is evidence that metrics are related to bird community composition (Mills et al., 1991; Francl & Schnell, 2002). Aspects of avian natural history may compound the anthropogenic/structural effects described above. In temperate regions, timing of migration (if the species migrates) and circadian activity patterns may affect the chance of BWCs. Kahle et al. (2016) found that BWCs increased during the periods of migration (April and October) and in mid-summer (July) when most birds are breeding. Numerous studies also concluded that BWCs were greatest during migration periods (Johnson & Hudson, 1976; Codoner, 1995; O’Connell, 2001; Gleb & Delacretaz, 2006; Hager et al., 2008). Despite the seasonal timing of these collisions, Klem (1989) concluded migratory status (as well as sex, age, and NO. 53, 2019 weather) did not affect BWCs. Supporting this, Blem & Willis (1998) and Kahle et al. (2016) concluded that migrating birds may not be major contributors to collisions. An examination of circadian activity patterns also presented us with conflicting data. While time of day was not a significant factor for BWCs by Klem (1989), others found clear trends in timing of collisions across daylight hours. Kahle et al. (2016) studied BWCs in an urban park (Golden Gate Park, San Francisco, California) and found that the greatest number of strikes occurred during mid-morning hours, 0900 — 1100 h. They found a steady decline in collisions throughout daylight hours. However, 83% of their documented BWCs occurred in daylight hours, as their pre-0900 h (early morning) surveys documented just 17% of the collisions. Other BWC studies concluded a priori that collisions were more likely during daylight hours, and limited their carcass surveys to afternoon time periods (e.g., 1400 — 1600 h; Hager & Consentino, 2014; Hager et al., 2017). In Virginia, BWCs have been investigated at a coastal campus at Old Dominion University (as part of a 40-campus national survey, individual results not presented in Hager et al., 2017) and in the western montane regions at the Virginia Tech Corporate Research Center (VTCRC) in Blacksburg. Although not the university campus proper, VIT'CRC does include 28 buildings (mostly 2-story, with maximum window areas of 693 m’) across 93 ha. In their study, they documented 240 bird casualties across 298 survey days. They discovered more BWCs with increased window area and an increase in ornamental vegetation around the buildings (Schneider et al., 2018). From this publication stemmed questions about nearby Radford University — a suburban campus set in a rural landscape — similar in land area to VT'CRC but with taller buildings at greater densities. Radford University, an 82.6-ha campus (37.13870°N, 80.55759°W; Fig. 1), is situated along a recognized migratory bird highway, the New River (e.g., VDGIF includes portions of the New River on its Bird & Wildlife Trail networks, VDGIF, 2019). Located about 20 km southwest of VT'CRC, this campus includes >30 buildings that are 3-4 stories tall, and one residence hall that is 13 stories tall. Construction on new and renovated buildings occurs year-round. Although the university seeks to build or renovate buildings so that they are LEED-certified, no buildings to date have incorporated bird-deterring windows or bird-deterring window decals (M. Biscotte, Office of Planning and Construction, Radford University). Furthermore, windows have been a substantial (window area > 1500 m*) component of facades at new or renovated buildings along major thoroughfares (Center for the Sciences, College of POWERS ET AL.: BIRD-WINDOW COLLISIONS A at” Xs Residency Gi = A Non-resident Te ZK s ane. < x Ch) A Resident ; o\ i “¥ LY og te Unidientified /* Cle 14 ) a es ~ : Jefferson > 2 & 4 S S Z S Ss “py & % O hey p Me Rec ~/\ >. — Center “yy AN - any S A 2 o- T N W ——" Moffett : 0 62.5 125 250, es || Kyle LS A\ ro Fig. 1. Radford University, Virginia campus map, with 15 buildings (labeled with names, in red) surveyed for bird-window collisions in 2018 and 2019. We recorded 0 — 8 hits/building. Locations of 51 documented BWC casualties, identified by species classified by residency status: non-residents (circle; N=15), residents (triangle; N=31), or unidentified (square; for birds not identified to species level, N=5). 13 14 BANISTERIA Humanities and Behavioral Sciences [CHBS] on Main street, Student Recreational Center on Jefferson St.; Fig. 2A, B, C). With these building additions and transformations in mind, we began a multi-year study to investigate BWCs at Radford University. Building on previous findings, we chose to investigate a number of potential landscaping or building features that could influence the location and number of BWCs: window area, building height, and two in situ vegetation metrics (total vegetation volume and the Levins index of vertical diversity). Next, we investigated features about the avian community: whether the birds were migratory (non-resident, transitory) or presumed resident species, and whether collisions likely occurred overnight or during daylight hours. We hypothesized that we would detect a greater number of BWCs at our newer buildings that possessed relatively greater window area, and that buildings with greater amounts and diversity of vegetation (which we perceive would reflect in the windows) would result in more BWCs. We further hypothesized that we would find no differences in BWCs between non-resident and resident species, and that most documented collisions would be discovered in the morning hours. METHODS Bird-window Surveys With the contributions from more than 30 Radford University students, we surveyed the perimeter of 15 campus buildings once or twice daily, ideally once in the morning and once in the afternoon. Buildings were selected to represent the full spectrum of building size (height and areal footprint), window area, and landscaped vegetation on campus. We completed surveys from 1 February 2018 through 15 November 2018, and from 7 February 2019 through 5 May 2019. From 6 May through 17 June 2019, we surveyed sporadically on 15 days. Following the protocol of Hager & Consentino (2014), surveyors walked within 2 m of building edges, scanning for potential bird hits; when a bird was discovered, photos were taken and its location was recorded in UTM. We classified legitimate hits as a cluster (>5) of feathers, partial body fragments, or whole bodies. Live, stunned birds also were also included as legitimate hits. We also collected carcasses opportunistically on campus, even if not collected at the 15 buildings and/or not during set surveys. For this reason, not all BWCs documented were included in every analysis. NO. 53, 2019 Bird Identification Participating students worked together to identify frozen full bird carcasses to species level, if possible, using standard bird field guides and museum specimens. For identification of partial carcasses and groups of feathers, we relied entirely on comparisons to preserved specimens from the Radford University Biology Department’s natural history collection (https://www.radford.edu/content/csat/home/biology/ facilities/natural-history-collection.html). Although not considered a valid BWC in this study, we collected and identified single feathers or small groups of <5 feathers, and retained them to build a library of known bird artifacts. If unidentifiable specimens contained tissue, they were examined via DNA barcoding analyses (see Paniagua-Ugarte et al., 2019). Landscape Analyses We (Powers) calculated total window area (m7) through analysis of architectural drawings of each of the 15 buildings. We (all authors) visited buildings and completed in-person measurements to confirm drawing specifications and remove from calculations windows that were opaquely painted. We determined maximum building height (m) through elevational metrics provided in the architectural drawings. Following methods similar to Francl & Schnell (2002), we measured vegetation in situ at points in ca. 40-m increments, around each building (5—18 points/building). We used a range pole, divided into seven 0.5-m increments (O—0.5 m, 0.5—1.0 m, ... 2.5—3.0 m, >3 m). We focused on vegetation at heights of 3 m or less because other studies reported that vegetation only affected BWCs at lower building floors (e.g., Gelb & Delatacruz, 2009). Standing ca. | m from the building facade, we documented a vegetation “hit” in the 0.5-m increment when vegetation was directly touching or within 10 cm of the pole. From these hits, we calculated two vegetation metrics: total vegetation volume (TVV; Mills et al., 1991) and the Levins index of vertical diversity (Levins, 1968). We estimated TVV using the formula: TVV =h/10v where h = number of intervals for which we documented vegetation hits, and v = total number of intervals (the number of points samples around the building). POWERS ET AL.: BIRD-WINDOW COLLISIONS 15 TER h eee PPrbay) BI > bldiahed LT re a => = Os Pa ae - ia tee ae Ie a ee ea Pees Sateiee TS Tee atl (| ' mee a } | lk =, Fig. 2. Examples of surveyed buildings at Radford University, documenting new (<5 years old) buildings on campus that incorporate large window areas (A: Center for the Sciences, B: College of Humanities and Behavioral Sciences [CHBS], C: Student Recreational Center) and more traditional buildings with lesser total window area (D: Whitt Hall, E: Trinkle Hall, F: Muse Hall). Photos by H. Reed, 2019. The Levins index is defined as: L =>. 1/ [(d;)"] 66599 1 where d; = total number of hits recorded for a 0.5-m increment total number of points measured around the building Statistical Analyses We utilized a forward stepwise regression comparing the number of BWC casualties per building to four metrics of each building: total window area, maximum building height, total vegetation volume, and Levins index. Setting a p-value of 0.25 to be included in the model, a priori, we ran the regression in JMP Pro 13 (SAS Institute, Cary, NC). We utilized a chi-square goodness of fit test in Microsoft Excel (Microsoft Excel 2019 MSO, Redmond, Washington) to determine if an equal number of carcasses were discovered in the morning (AM) versus the evening (PM) surveys. We considered morning hits as those discovered in daylight surveys completed from ca. 0600—1200 h. Evening hits were those discovered from 1201—1800 h. We recognize that hits that occurred overnight (1801 h—0559 h) are lumped with the morning collections, and we would therefore expect that, if collision patterns were random, we would expect 75% of carcasses to be collected during morning surveys and 25% during evening surveys. Further, we limited our analyses to birds collected on dates in which two surveys/day were completed, so that we could confidently assign the correct collision time block. In the same manner, we used a chi-squared goodness of fit test in Microsoft Excel to determine if an equal number of hits occurred for birds considered residents versus those actively migrating (i.e., suspected to collide with buildings while in novel surroundings). Here, we defined “resident” as a bird who is present in the area year-round or migratory but a full- time inhabitant during summer months. These birds would be expected to be familiar with the surroundings. Birds were assigned migratory, non-resident status if they were collected during the species’ known migration period; we assumed the area was unfamiliar to them. Residency status was derived from Cornell Lab of Ornithology’s Birds of North America (Rodewald [Ed.], 2015) using geographic range maps, text, and annual cycle figures (when available). For questionable birds whose migratory status was unclear in southwestern Virginia, we further investigated status utilizing information from the Virginia Breeding Bird Atlas (https://ebird.org/atlasva) and Christmas Bird Counts (https://www.audubon.org/conservation/science/ christmas-bird-count) from the region. Migratory status could only be assigned for ca. six months of the year (3 months for spring migrations, three months for fall migrations) and resident status could be assigned year- round. Therefore, we expected to detect twice as many residents as non-residents by chance alone. 16 BANISTERIA RESULTS In >975 hours of surveys across 393 days, we documented 51 BWCs across 23 species at Radford University (Appendix 1; Figure 1, 3). BWCs/building ranged from 0-8. Of the 51 birds, one American robin (Turdus migratorius) was founded alive but stunned (Fig. 3D); it flew away when the observer attempted to collect it. Sixteen full carcasses (deceased, Fig. 3A, B, C) and 34 partial carcasses or piles of feathers also were collected (Fig. 3E, F). Fifteen individuals across 10 Species were non-residents. We found that 31 individuals among 15 species were resident species. Five songbird individuals were not able to be fully identified to the species level, and were not included in this analysis (Fig. 1). A chi-squared goodness-of-fit test for these 46 individuals revealed no significant difference between resident and non-resident species BWC rates (y7=0.011, df=1, p=0.917). In the 392 days of surveys, we completed 863 individual campus walks. In 827 walks in which time was recorded, 368 were completed in the morning time block and 459 were evening surveys. When we factored out days in which single walks were completed (1.¢e., we were not able to confidently determine which time block the collision actually occurred), our sample size was reduced to 18 documented collisions. Recording ten hits in the AM block and eight in the PM block, our chi- squared goodness-of-fit test suggested that PM hits occurred marginally more than expected by chance alone (y?=0.363, df=1, p=0.056). Forty-eight of our 51 BWCs occurred at the 15 buildings for which we calculated window area, building height, and quantified vegetation. Our forward stepwise regression, comparing the number of hits per building versus the four variables reported that the only significant variable was window area (1? = 0.335, F = 6.558, p=0.024: Table 1). As window area increased, so did the number of BWCs/building. NO. 53, 2019 DISCUSSION Our finding that window area was the only metric significantly affecting BWC was not surprising, as the majority of BWC studies have detected this same primary factor across the landscape. Building height may have been less of a factor on this campus because, as originally stated, most buildings are of similar height; however, newer constructions and _ renovations incorporate markedly more windows into their facades. Perhaps time since construction may have been a co-predictor (with window area) of BWCs on campus, but this metric may not be transferrable to other studies. We also failed to find any vegetation effects on BWCs. Because of the plethora of metrics utilized to measure these features, we either selected metrics that did not accurately account for vegetation around buildings, or we looked at too fine of a scale for vegetation to have affected these birds. Perhaps future studies will rely instead on a broad-scale GIS component, as several studies did find significant, though secondary, effects of vegetation on BWCs (e.g., Hager et al., 2017; Schneider et al., 2018). The discovery of only 51 bird carcasses in nearly 1,000 h and 393 days of surveys is surprisingly low, compared to other BWC studies across the continent. Locally, Schneider et al. (2018) documented 240 individuals in a shorter time span, only surveying “when schedules and weather allowed.” Our efforts were highest (2 surveys/day) during the fall and spring semesters, which should have corresponded with migratory patterns of birds. We were, at the very least, consistent (1 survey/day) during summer months and when school was not in session. Our efforts attempted to minimize time for scavengers to access the carcasses, yet only 16 (17 if the stunned, live bird is included) full, intact carcasses were discovered. The remaining 34 birds suggested scavenging had occurred (Fig. 3E, F). Nocturnal scavenging events would be expected, as Table 1. Results of forward stepwise regression, examining factors influencing number of bird-window collisions per building at 15 buildings on Radford University’s campus, 2018-2019. Of four metrics, window area alone explained 33.5% of total variance; no other variables were included in the final model. Parameter Estimate DF Intercept 1.6770 ] Window area (m7) 0.0017 1 Building height (max., m) 0 1 Levins 0 1 TVV 0 1 SS F p 0) 0) 1.000 44.576 6.558 0.023 1.537 0.212 0.653 1.231 0.170 0.688 7939 1.119 0.311 POWERS ET AL.: BIRD-WINDOW COLLISIONS 17 Fig. 3. Examples of casualties from bird-window collisions. Of the 51 documented collisions, 16 were whole bodies - deceased (e.g., A: House Finch [Haemorhous mexicanus], B: Chimney Swift [Chaetura pelagica], C: Yellow-billed Cuckoo [Coccyzus americanus]), 1 was stunned but recovered (D: American Robin [Turdus migratorius]), and the rest were portions of scavenged carcasses (e.g., E: Gray Catbird [Dumetella carolinensis], F: White-breasted Nuthatch [Sitta carolinensis]). Photos by six participating students at Radford University, 2018-2019. personal observations include Striped Skunks (Mephitis mephitis), Raccoons (Procyon lotor), Virginia Opossums (Didelphis virginiana), and feral cats as on- campus visitors. Future studies may involve setting a wildlife camera on planted carcasses to determine frequency of and time until documented scavenging or unanticipated anthropogenic disturbances, like students or facilities workers collecting the carcass. As multiple studies have acknowledged, it is likely that our 51 mortalities are underreporting the actual number of BWCs (Bayne et al., 2012; Kummer et al., 2016b). Besides carcass scavenging (Hager et al., 2012), observer bias plays a significant role in documenting BWCs. With over 30 (albeit trained) students contributing to our project, we assume the visual acuity, mental focus, and ability to detect feathers and partial or full carcasses varied by student (Hager & Cosentino, 2014). In other bird carcass surveys, researchers suggest that the actual number of bird mortalities is 2.3—5 times greater than what is discovered (Dunn, 1993; Zimmerling et al., 2013). Furthermore, despite the finding from other studies (e.g., Gelb & Delacretaz, 2006; Kahle et al., 2016) that most BWCs occurred during daylight hours, and a marginally significant finding to support that, we are not confident about our sample size. Our intensive twice-daily surveying efforts were too inconsistent across the study, and we could only include 18 of the 51 carcasses for statistical analysis. Our future efforts on campus may investigate short (2-3 week) efforts at buildings with the highest rate of collisions. We might complete three surveys daily, at 8-h intervals, to tease apart collision-time trends. The shorter time frame and subset of buildings might make such studies temporally feasible, given student schedules. The near-absence of rare or protected species in our observations is interesting. Indeed, none of the 23 species are listed as species of greatest conservation need in the Virginia Wildlife Action Plan (VDGIF, 2015), and none are afforded state-threatened or endangered status. Although we documented three fairly uncommon warblers — Magnolia (Setophaga magnolia, 10 October 18 BANISTERIA 2018; S2B status — suggesting they are rare breeders in Virginia, Wilson & Tuberville, 2003), Cape May (S. tigrina;, 29 September 2018), Worm-eating (He/mitheros vermivorum, 7 May 2018) - all were collected during peak migration periods. Indeed, eBird records document other individuals in the area — some along the New River in Radford — within a two-week window of these finds (https://ebird. org/atlasva/explore). Of the 23 species documented, only the Swainson’s Thrush (Catharus ustulatus) was an unexpected seasonal find. Documented on 30 June 2018, the timing is long after the putative migratory season has concluded; the only Radford record of this species on eBird was on 23 May 2016 (reported by C. Kessler, https://ebird.org/ atlasva/map/swathr), coinciding with migration periods. Furthermore, just one June record has been reported from nearby counties (Giles Co., VA/Monroe Co., WV line, C. Kessler, pers. comm.). Our mid-summer collision record suggests this individual may have been maintaining a summer residence in the area. This species was identified only by DNA analyses (Paniagua-Ugarte et al., 2019), and we cannot know the age, sex, or any other natural history characteristics of this individual. This species is state- ranked as S1B (Wilson & Tuberville, 2003), suggesting that it is an extremely rare breeder in_ the Commonwealth. The Virginia Fish and Wildlife Information Service (VaFWIS) system also indicates that this species has not been documented in Radford City in June and that all regional records of Swainson’s Thrush were reported during the migratory seasons (S. Watson, VDGIF, pers. comm.). The natural history of this species in our region certainly warrants further investigation. Twenty-one of our 51 BWCs occurred at only three buildings, all newly-constructed in the last five years (Fig. 2A, B, C) and all possessing substantial window areas (1685-3865 m’). As it appears that Radford University is implementing greater window areas in new construction, we strongly suggest that bird- deterring efforts be applied. Window decals can be useful on a small-scale, and even applied on a window- by-window basis by concerned faculty members (as many personal offices contain windows). However, it is unlikely that decals, typically with patterns to make the window more visible to the birds, could or would be utilized on aspects whose window areas comprise nearly 100% of the facade (e.g., Fig. 2A, B, C). The American Bird Conservancy has published a number of window types and the related “threat factor’ for BWCs (American Bird Conservancy, 2012). Patterned glass (simple, vertical lines are suggested), translucent glass, and glass coated with UV-reflecting lines all could reduce BWCs, and contribute to LEED-certification (Klem, 2009; Green Building Alliance, 2016). Currently, NO. 53, 2019 bird-deterring window modifications offer a pilot credit towards said certification (American Bird Conservancy, 2012). Our project, therefore, provides useful information to the Radford University Office of Planning and Construction, as they design and implement the renovations and new construction on campus. Our data will help the university identify existing areas for potential treatment, as funds become available (M. Biscotte, Radford University Office of Planning and Construction). Implementing such building modifications could establish Radford University as a leader in “green” architecture and provide new research opportunities for students in coming years. ACKNOWLEDGEMENTS We are grateful to the more than 30 Radford University students who contributed to BWC surveys. We thank Michael Biscotte and Benny Skeens of the Radford University Office of Planning and Construction providing historical and contemporary architectural drawings of the campus buildings. The Radford University Department of Biology supported this research-based course in bird-window collisions. LITERATURE CITED American Bird Conservancy. 2012. Bird-friendly building design. http://collisions.abcbirds.org/ (Accessed 29 June 2019). Bayne, E. M., C. A. Scobie, & M. Rawson-Clark. 2012. Factors influencing the annual risk of bird window collisions at residential structures in Alberta, Canada. Wildlife Research 39: 583-592. Blem, C. R., & B. A. Willis. 1998. Seasonal variation of human-caused mortality of birds in the Richmond area. Raven 69: 3-8. Briscoe, T., & C. Dampier. 2019. As many as a billion birds are killed crashing into buildings each year - and Chicago’s skyline is the most dangerous area in the country. Chicago Tribune, 4 April. Chin, S. 2016. Investigating the effects of urban features on bird window collisions. M.S. thesis, York University, Toronto, Ontario, Canada. 46 pp. Codoner, N. A. 1995. Mortality of Connecticut birds on roads and at buildings. Connecticut Warbler 15: 89-98. Dunn, E. H. residential windows 1993. Bird mortality from. striking in winter. Journal of Field POWERS ET AL.: BIRD-WINDOW COLLISIONS 19 Ornithology 64: 302-309. Francl, K. E., & G. D. Schnell. 2002. Relationships of human disturbance, bird communities, and plant communities along the land-water interface of a large reservoir. Environmental Monitoring and Assessment 73: 67-93. Gelb, Y., & N. Delacretaz. 2006. Avian window strike mortality at an urban office building. Kingbird 56: 190— 198. Gelb, Y., & N. Delacretaz. 2009. Windows and vegetation: Primary factors in Manhattan bird collisions. Northeastern Naturalist 16: 455-470. Green Building Alliance. 2016. Bird-friendly design. https://www.go-gba.org/resources/green-building- methods/bird-friendly-design/ (Accessed 16 May 2019). Hager, S. B., & B. J. Cosentino. 2014. Surveying for bird carcasses resulting from window collisions: a standardized protocol. PeerJ PrePrints 2:e406v1 https://doi.org/10.7287/peerj.preprints.406v1 Hager, S. B., B. J. Cosentino, & K. J. McKay. 2012. Scavenging affects persistence of avian carcasses resulting from window collisions in an urban landscape. Journal of Field Ornithology 83: 203-211. Hager, S. B., B. J. Cosentino, M. A. Aguilar-Gomez, & 52 others. 2017. Continent-wide analysis of how urbanization affects bird-window collision mortality in North America. Biological Conservation 212(A): 209— 2A Hager, S. B., H. Trudell, K. J. McKay, S. M. Crandall, & L. Mayer. 2008. Bird density and mortality at windows. Wilson Journal of Ornithology 120: 550-564. Johnson, R. E., & G. E. Hudson. 1976. Bird mortality at a glassed-in walkway in Washington State. Western Birds 7: 99-107. Kahle, L. Q., M. E. Flannery, & J. P. Dumbacher. 2016. Bird-window collisions at a west-coast urban park museum: Analyses of bird biology and window attributes from Golden Gate Park, San Francisco. PloS ONE 11(1): e0144600. doi:10.1371/journal.pone.0144600 Klem Jr., D. 1989. Bird-window collisions. Wilson Bulletin 101: 606-620. Klem Jr., D. 2009. Preventing bird-window collisions. Wilson Journal of Ornithology 121: 314-321. Kummer, J. A., E. M. Bayne, & C. S. Machtans. 2016a. Use of citizen science to identify factors affecting bird— window collision risk at houses Condor 118: 624-639. Kummer, J. A., C. J. Nordell, T. M. Berry, C. V. Collins, C.R. L. Tse, & E. M. Bayne. 2016b. Use of bird carcass removals by urban scavengers to adjust bird-window collision estimates. Avian Conservation and Ecology 11(2): 12. _ http://dx.doi.org/10.5751/ACE-00927- 110212 Leighton, G. M., J. H. Lee, & K. E. Francl. 2009. Influence of structural complexity on bat activity at palustrine habitats in the northern Great Lakes region. Michigan Academician 39: 33-46. Levins, R. 1968. Evolution in Changing Environments: Some Theoretical Explorations, Monographs in Population Biology, No. 2, Princeton University Press, Princeton, NJ. 132 pp. Loss, S. R., T. Will, S. S. Loss, & P. P. Marra. 2014. Bird-building collisions in the United States: Estimates of annual mortality and species vulnerability. Condor 116: 8-23. Mills, G. S., J. B. Dunning, & J. M. Bates. 1991. The relationship between breeding bird density and vegetation volume. Wilson Bulletin 103: 468-479. O’Connell, T. J. 2001. Avian window strike mortality at a suburban office park. Raven 72: 141-149. Paniagua-Ugarte, C. Y., K. E. Powers, & R. R. Sheehy. 2019. Using DNA barcoding to identify carcasses from bird-window collisions at Radford University. Banisteria 53: 22-26. Schneider, R., C. Barton, K. Zirkle, C. Greene, & K. Newman. 2018. Year-round monitoring reveals prevalence of fatal bird-window collisions at the Virginia Tech Corporate Research Center. PeerJ e4562. https://peerj.com/articles/4562/ The Birds of North America (P. Rodewald, ed.). Ithaca, NY: Cornell Laboratory of Ornithology; Retrieved from The Birds of North America: https://birdsna.org; AUG 2015 (Accessed 20 August 2019). Virginia Department of Game and Inland Fisheries 20 BANISTERIA (VDGIF). 2015. Virginia Wildlife Action Plan. http://bewildvirginia.org/wildlife-action-plan/ (Accessed 30 June 2019). Virginia Department of Game and Inland Fisheries (VDGIF). 2019. Virginia Bird and Wildlife Trails. https://www.degif. virginia. gov/vbwt/ (Accessed 14 May 2019). Wilson, I. T., & T. Tuberville. 2003. Virginia’s Precious Heritage: A Report on the Status of Virginia’s Natural Communities, Plants, and Animals, and a Plan for NO. 53, 2019 Preserving Virginia’s Natural Heritage Resources. Natural Heritage Technical Report 03-15. Virginia Department of Conservation and Recreation, Division of Natural Heritage, Richmond, VA. 82 pp. plus appendices. Zimmerling, J. R., A. C. Pomeroy, M. V. d’Entremont, & C. M. Francis. 2013. Canadian estimate of bird mortality due to collisions and direct habitat loss associated with wind turbine developments. Avian Conservation and Ecology 8(2): 10. http://dx. doi.org/10.5751/ace-00609-080210 POWERS ET AL.: BIRD-WINDOW COLLISIONS 21 Appendix 1. List of 51 individual birds across 23 confirmed species that were BWC casualties at Radford University in 2018 (February—October) and/or 2019 (February—June). An “*” indicates that one individual was confirmed via genetic analyses of carcass tissue (Paniagua-Ugarte et al., 2019). Family Order Caprimulgiformes Apodidae Order Columbiformes Columbidae Columbidae Order Cuculiformes Cuculidae Order Passeriformes Bombycillidae Cardinalidae Certhiidae Corvidae Fringillidae Fringillidae Icteridae Mimidae Paridae Parulidae Parulidae Parulidae Passerellidae Passerellidae Passeridae Sittidae Turdidae Turdidae Turdidae Scientific name Chaetura pelagica Columba livia Zenaida macroura Coccyzus americanus Bombycilla cedrorum Cardinalis cardinalis Certhia americana Corvus brachyrhynchos Haemorhous mexicanus Spinus tristis Molothrus ater Dumetella carolinensis Baeolophus bicolor Helmitheros vermivorum Setophaga magnolia Setophaga tigrina Melospiza melodia Spizella passerina Passer domesticus Sitta carolinensis Catharus ustulatus Sialia sialis Turdus migratorius Common name Chimney Swift Rock Dove Mourning Dove Yellow-billed Cuckoo Cedar Waxwing* Northern Cardinal Brown Creeper American Crow House Finch American Goldfinch Brown-headed Cowbird Gray Catbird Tufted Titmouse Worm-eating Warbler Magnolia Warbler Cape May Warbler* Song Sparrow Chipping Sparrow House Sparrow* White-breasted Nuthatch Swainson’s Thrush* Eastern Bluebird American Robin Unknown songbird Number of individuals Banisteria, Number 53, pages 22—26 © 2019 Virginia Natural History Society Using DNA Barcoding to Identify Carcasses from Bird-window Collisions at Radford University Claudia Y. Paniagua-Ugarte, Karen E. Powers', and Robert R. Sheehy Biology Department Radford University Radford, Virginia 24142 ‘Corresponding author: kpowers4@radford.edu ABSTRACT A leading cause of avian mortality is collisions with building windows. To fully understand the impacts of bird- window collisions at Radford University, Virginia, bird carcasses (whole or in part) were collected and documented in 2018-2019. Although the majority of carcasses were identified via morphological features, the taxonomic identification of some samples was impossible due to evidence of predation, environmental degradation, and confusion in species differentiation due to sex, age, and seasonal plumage. We used DNA barcoding to identify carcasses in cases where species identification based on morphology was not possible. DNA barcoding with standard PCR primers allowed for the successful identification of five individuals across four species - two of which species had not been previously identified in this study. Our study emphasizes the application of DNA barcoding in bird-window collision studies, and its potential for use in other conservation and mitigation efforts. Keywords: avian mortality, cytochrome c oxidase subunit 1, species identification, taxonomy. INTRODUCTION Bird-window collisions (BWCs) are a leading cause of mortality in the United States and word-wide, with annual mortality estimates of nearly one billion birds (Loss et al., 2014). Such collisions are suspected to be greater during peak migratory periods, as birds traverse less familiar habitats (Borden et al., 2010; Schneider et al., 2018). Borden et al. (2010) reported that migratory birds were nine times more likely to be a casualty of BWCs than resident species. While some studies report that particular species or family groups are more prone to collisions (e.g., hummingbirds, Schneider et al., 2018), it is evident that both common and rare species are susceptible to this threat. Multiple large-scale and/or long-term studies of BWCs acknowledge that mortalities are likely under- reported (Bayne et al., 2012; Kummer et al., 2016). A major limitation is observer error, which manifests itself as a failure to detect carcasses that are present (e.g., carcasses obscured by vegetation). Carcass scavenging is a second major cause of BWC underestimates (e.g., Kummer et al., 2016). Finally, carcasses landing outside a limited search area may also affect discovery (Zimmerling et al., 2013). Recent estimates suggest that carcass recovery is 2.3—5 times less than the actual number of bird mortalities (Dunn, 1993; Zimmerling et al., 2013). Identification is important because all bird samples collectively play a role in understanding how species migration, seasonal distribution, and density could relate to bird building collisions (Schneider et al., 2018). However, once recovered, several circumstances may limit accurate identification of the carcass. Damage to the specimen resulting from the collision, length of time between mortality and collection, and scavenger activity may all affect the ability to identify that carcass. Additionally, differentiation between juvenile and adult females may be difficult; particularly among passerines. In our BWC study (Powers et al., this volume), all of these factors impeded accurate identification of some specimens. From February 2018 — June 2019, students at Radford University investigated myriad aspects of natural and anthropogenic influences on the number and nature of BWCs at 15 buildings on the university campus (Powers et al., this volume). Statistical analyses required the accurate identification of bird carcasses. Students PANIAGUA-UGARTE ET AL.: USING DNA BARCODING 23 identified full carcasses, partial carcasses, and multiple feather evidence from BWCs via comparisons to bird specimens within the natural history collection at Radford University. However, morphological identification of nine of 51 birds was not possible or definitive. DNA barcoding, identifying species by comparing a short, defined DNA sequence to a DNA sequence reference database, provided an alternative approach to identifying a specimen (Kerr et al., 2007). DNA barcoding has been effective for the species identification of whole birds (Herbert et al., 2004), bird tissue (Dove et al., 2008), and eggs (Lee & Prys-Jones, 2008) from unknown samples. Of the nine unidentifiable carcasses, seven contained tissue, and thus had the potential to be identified through DNA barcoding. Our goal was to extract DNA from carcasses and use DNA barcoding to identify them to the fullest extent possible. METHODS Seven bird carcasses that contained tissue but were physically unidentifiable were collected and stored in a standard chest freezer at 20° C. One additional BWC specimen collected in October 2016 at Radford University was included in this project, increasing our sample size to eight. DNA was extracted from tissues using the Qiagen DNeasy blood and tissue kit (Qiagen, Austin, Texas). We amplified a 708 bp fragment of the mitochondrial encoded cytochrome c oxidase subunit I (COI) gene using tailed primers BirdFl tl and BirdR1 tl (Kerr et al., 2007). Each 50 ul PCR reaction contained 25 ul of 2x Quick-Load Taq master mix (NEB, Ipswich, Massachusetts) 2.3 mM MgClo, 0.5 mM each of forward and reverse primers, 5 ul of DNA template (approx. 200 ng), and distilled water. PCR amplification was performed with an initial denaturation (3 min at 94° C) followed by 35 cycles of 94° C for 1 min, 51° C for 1 min, and 68°C for 1 min. This was followed by a final extension step at 68° C for 10 min. PCR reactions were held at 4° C for 6 to 8 h until storage at -20° C. We identified successful amplification by running 5 ul of each sample on 2% agarose/TBE/EtBr gel. Successfully amplified samples were sent to GENEWIZ (South Plainfield, New Jersey) for sequencing. Both strands of our amplicons were sequenced using the Sanger dideoxy chain termination method (Sanger & Coulson, 1975). We used the DNA subway (DNA Learning Center, https://dnasubway.cyverse.org) as a means to align forward and reverse sequences of each sample and to evaluate/correct discrepancies between the sequences of the forward and reverse strands and to construct a consensus sequence of the forward and reverse strands. We used the “Identification Engine” (http://boldsystems.org/index.php/IDS_OpenIdEngine) to identify the taxonomic origin of unknown specimen DNA using the consensus sequence as a query. We assume “correct” species identification for samples that show 299.0% sequence similarity with bird species already in the database. RESULTS Of the eight carcasses tested, we successfully amplified and identified five to species level (Table 1). We confidently identified one Cedar Waxwing (Bombycilla cedrorum), one House Sparrow (Passer domesticus), two Cape May Warblers (Setophaga tigrina, one was the 2016 collection), and one Swainson’s Thrush (Catharus ustulatus). The thrush and the Cape May Warblers had not been previously identified in the BWC project. The specimens whose DNA did not amplify remain unidentified via morphological features. DISCUSSION The use of DNA barcoding allowed for the identification of five of the eight unidentifiable bird carcasses. All five of these carcasses had a sequence similarity of =99.0% to reference species and are considered to be accurately identified. Unsuccessful amplification may be due to the degradation of DNA that results with age and/or the environmental conditions experienced before collection. There are, however, various methods that may allow for the analysis of degraded DNA such as the PCR amplification of shorter sequences using internal primers. While we used primers designed to amplify the complete 708 bp “Folmer region” of the COI gene (Folmer et al., 1994), these primers prove ineffective at amplifying all samples. Often, this results from the fragmentation of DNA due to environmental degradation which results in the lack of a full-length DNA template. Future research will be aimed at designing internal primers which will allow the amplification of shorter target sequences and/or the application of DNA mini- barcodes (Meusnier et al., 2008) to these problematic specimens. This should allow us to apply DNA barcoding to a more degraded template. The results from the five successfully identified samples provide useful data to help us understand the relationships of particular bird species with BWCs. Of greatest interest were the two species (Swainson’s Thrush and Cape May Warbler) that had not previously been identified as BWC casualties in this project. Both are migratory species in the region and not considered 24 BANISTERIA NO. 53, 2019 Table 1. Bird-window collision carcasses collected from Radford University (2016, 2018-2019) that were identified with DNA barcoding. Presented is a photo of each original carcass, the range of sequence similarity to members of the top matched species, and the DNA percent match, broken down by family, genus, and specific epithet. All samples fall into the order Passeriformes. SAMPLE TAXON PROBABILITY OF SEQUENCE SIMILARITY ASSIGNMENT PLACEMENT TO TOP 100 DATABASE (%) MATCHES Bombycillidae 100 nt Bombycilla 100 é i cedrorum 100 z i Cedar Waxwing vs 12 2 uM Passeridae 100 ‘a98 Passer 100 e oe domesticus 100 i m9 House Sparrow ae ——— Parulidae 100 way Setophaga 100 aa tigrina 99.7 : te Cape May Warbler al —— Parulidae 100 ‘os Setophaga 100 : sao tigrina 100 ; “ Cape May Warbler m5 2 2 «4s Turdidae 100 por Catharus 100 c om ustulatus 100 i vs Swainson’s Thrush seo. PANIAGUA-UGARTE ET AL.: USING DNA BARCODING 25 summer residents. Radford University’s campus abuts the New River, a migratory pathway. Therefore, it is not entirely unexpected to find these species. While both species are uncommon, they have been documented along the New River in Radford. The Cape May Warbler has been observed in late September and early October, and the Swainson’s Thrush has been observed most often in September through mid-October with sporadic observations in May (e.g., eBird data for Bisset Park and Riverway Trail, Radford, Virginia; https://ebird.org/atlasva). The Swainson’s Thrush was collected on 30 June 2018, providing the first local record of this taxon outside regular migration periods. To date, only one June record in the region has been reported (Giles Co., VA/Monroe Co., WV line, C. Kessler, pers. comm.). Powers et al. (2019) review these species in greater detail. The identification of the Swainson’s Thrush demonstrates the power of DNA barcoding as this potentially significant observation would have otherwise gone unnoticed. We have demonstrated the power of DNA barcoding in providing an alternative means for the taxonomic identification of specimens where, for numerous and varied reasons, traditional means of identification based on morphology may be inconclusive or impossible. While we have applied this approach to the identification of avian taxa, it is easy to see where this approach would be useful in other studies as well. DNA barcoding could easily be applied to the taxonomic identification of tissues from roadkills, prey remains found in association with predatory animals, and cryptic species where the identification based on morphological differences is problematic. These methods could be particularly useful in regulatory projects, like surveys around wind turbines (for birds and bats), where identification to species level may have greater implications for mitigation efforts. Applying DNA barcoding as an approach for species identification should not fall outside the realm of many studies. With the exception of DNA sequencing, the molecular techniques necessary to complete this project are common among introductory biology and genetics college-level courses. Myriad of companies are available to perform the Sanger dideoxy sequencing reaction if these are unavailable in-house. The cost per sample of DNA barcoding can range from between $5.00 to $10.00 USD depending upon: in-house versus commercial sequencing, sequencing one or both strands or, the necessity for band isolation to remove primer dimers. If one is distinguishing among only a few potential taxa, a related approach, Cleaved Amplified Polymorphic Sequences (CAPS; Konieczny & Ausubel, 1993) or the related dCAPS (derived Cleaved Amplified Polymorphic Sequences; Neff et al., 1998) could successfully be applied to species identification saving time and reducing the cost of analysis to a few dollars. ACKNOWLEDGEMENTS We thank the Department of Biology at Radford University for covering laboratory expenses associated with this bird-window collision research, and for supporting two courses in this research. We thank the more than 30 students at Radford University who contributed nearly 1,000 hours of surveys in this bird- window collision project. LITERATURE CITED Bayne, E. M., C. A. Scobie, & M. Rawson-Clark. 2012. Factors influencing the annual risk of bird-window collisions at residential structures in Alberta, Canada. Wildlife Research 39: 583-592. Borden, W. C., O. M. Lockhart. A. W. Jones, & M. S. Lyons. 2010. Seasonal, taxonomic, and local habitat components of bird-window collisions on an urban university campus in Cleveland, OH. Ohio Journal of Science 110: 44—52. Dove, C. J., N. C. Rotzel, M. Heacker, & L. A. Weigt. 2008. Using DNA _ barcodes to identify bird species involved in birdstrikes. Journal of Wildlife Management 72: 1231-1236. Dunn, E. H. 1993. Bird mortality from. striking residential windows in winter. Journal of Field Ornithology 64: 302-309. Folmer, O., M. Black, W. Hoeh, R. Lutz, & R. Vrijenhoek. 1994. DNA primers for amplification of mitochondrial cytochrome c oxidase subunit I from diverse metazoan invertebrates. Molecular Marine Biology and Biotechnology 3: 294-299. Herbert, P. D., M. Y. Stoeckle, T. S. Zemlak, & C. M. Francis. 2004. Identification of birds through DNA barcodes. PLoS Biology 2(10): e312. https://doi.org/10.1371/journal.pbio.0020312 Kerr, K. C., M. Y. Stoecke, C. J. Dove, L. A. Weight, C. M. Francis, & P. D. Hebert. 2007. Comprehensive DNA barcode coverage of North American birds. Molecular Ecology Notes 7: 535-543. Konieczny A., & F. M. Ausubel. 1993. A procedure for mapping Arabidopsis mutations using co-dominant 26 BANISTERIA ecotype-specific PCR-based markers. Plant Journal 4: 403-410. Kummer, J. A., C. J. Nordell, T. M. Berry, C. V. Collins, C.R. L. Tse, & E. M. Bayne. 2016. Use of bird carcass removals by urban scavengers to adjust bird-window collision estimates. Avian Conservation and Ecology 11(2): 12. http://dx.doi.org/10.575 1/ACE-00927-11021. Lee, P. L., & R. P. Prys-Jones. 2008. Extracting DNA from museum bird eggs, and whole genome amplification of archive DNA. Molecular Ecology Resources 8: 551—560. Loss, S. R., T. Will, S. S. Loss, & P. P. Marra. 2014. Bird-building collisions in the United States: Estimates of annual mortality and species vulnerability. Condor 116: 8-23, Meusnier, I., G. A. C. Singer, J. F. Landry, D. A. Hickey, P. D. N. Hebert, & M. Hajibabaei. 2008. A universal DNA mini-barcode for biodiversity analysis. BMC Genomics 9: 214-218. Neff, M. M., J. D. Neff, J. Chory, & A. E. Pepper. 1998. dCAPS, a simple technique for the genetic analysis NO. 53, 2019 of single nucleotide polymorphisms: Experimental applications in Arabidopsis thaliana genetics. Plant Journal 14: 387-392. Powers, K.E., L.A. Burroughs, B.M. Mullen, H.C. Reed, & Z.Q. Krajcirovic. 2019. Investigating campus features that influence bird-window collisions at Radford University, Virginia. Banisteria 53: 11-21. Sanger, F., & A. R. Coulson. 1975. A rapid method for determining sequences in DNA by primed synthesis with DNA polymerase. Journal of Molecular Biology 94: 441-448. Schneider, R. M., C. M. Barton, K. W. Zirkle, C. F. Greene, & K. B. Newman. 2018. Year-round monitoring reveals prevalence of fatal bird-window collisions at the Virginia Tech Corporate Research Center. PeerJ e4562. https://peerj.com/articles/4562/ Zimmerling, J. R., A. C. Pomeroy, M. V. d’Entremont, & C. M. Francis. 2013. Canadian estimate of bird mortality due to collisions and direct habitat loss associated with wind turbine developments. Avian Conservation and Ecology 8(2): 10. http://dx.doi.org/10.575 1/ACE-00609-080210 Banisteria, Number 53, pages 27—71 © 2019 Virginia Natural History Society The Rove Beetles (Coleoptera: Staphylinidae) of the George Washington Memorial Parkway, with a Checklist of Regional Species R. Michael Brattain 505 Lingle Terrace Lafayette, Indiana 47901 Brent W. Steury! U.S. National Park Service 700 George Washington Memorial Parkway Turkey Run Park Headquarters McLean, Virginia 22101 Alfred F. Newton and Margaret K. Thayer Field Museum of Natural History 1400 South Lake Shore Drive Chicago, Illinois 60605 Jeffrey D. Holland Department of Entomology Purdue University 901 West State Street West Lafayette, Indiana 47907 ABSTRACT Two-hundred and nine taxa (171 identified to species level), in 111 genera, of staphylinid beetles were documented during a 21-year field survey of a national park site (George Washington Memorial Parkway) that spans parts of Fairfax and Arlington counties in Virginia. Fifty-two species, plus five additional genera, documented from the Parkway are first records for Virginia. An additional 62 species new to Virginia are listed in the appendix from broader research documenting 792 species of Staphylinidae from Virginia, Maryland, and the District of Columbia. The study also increases the number of staphylinid species known from the Potomac River Gorge to 167. Of the seven capture methods used in the survey, Malaise traps were the most successful. 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. Notes on morphological characteristics and habitats are given for some species. Thirteen species adventive to North America are documented from the Parkway and 60 adventive species are recorded from Virginia, Maryland, and the District of Columbia in the Appendix. Range extensions are documented for 16 species. Images of 11 species new to Virginia are provided. Keywords: Biodiversity, District of Columbia, insects, Malaise traps, Maryland, national parks, new state records, Potomac River Gorge, Virginia. ‘Corresponding author: brent_steury@nps.gov 28 BANISTERIA INTRODUCTION Probably the largest family of beetles in the world, the Staphylinidae, or rove beetles, contains more than 64,031 species in 32 subfamilies and 167 tribes (Irmler et al., 2018; Newton, 2019). It is the largest beetle family in North America, with 568 genera containing over 4,500 species (Newton et al., 2000; Newton, 2019). They are generally recognized by their short, truncate elytra that leave exposed a large dorsal part of the abdomen. In many tribes the abdomen is flexible and is waggled from side to side as the beetle moves along the ground. In a few groups the abdomen is less flexible (Micropeplinae) or the elytra cover the abdomen (some Omaliinae, Scydmaeninae). Possible staphylinid fossils have been found near the Jurassic/Triassic boundary that are more than 200 million years old, and numerous subfamilies were present by the end of the Jurassic (Newton, et al., 2000; Chatzimanolis, 2018). No other group of Coleoptera has been as successful as the Staphylinidae at living in such an enormous number of diverse habitats (Thayer, 2016; Betz et al., 2018). Adults of most species are nocturnal or shelter in dark areas during daylight hours; some exceptions include Stenus species, which are active in daylight. They are common components of soil biota, found in decaying leaf litter or deeper soil layers where they prey upon a variety of soil inhabiting organisms, or feed on decaying plant material or on fungi. However, this family fills nearly every ecological niche and they can be found in fungi, rotting wood, dung, carrion, caves, mammal burrows, and ant or termite nests. Some smaller species (most species are 2-8 mm) have an ant-like habitus and some species (especially Aleocharinae) live among particular ants or other social Hymenoptera or termites. Others are associated with birds or ectodermally on mammals (Brunke & Buffam, 2018). These associations can be beneficial (consuming the host waste material) or parasitic, when they prey on the eggs, larvae or stored food of the hosts. The larvae of Aleochara are parasitoids of dipterous puparia. Because of their hyper-diversity of form, habitats, and prey, their ancient origins, and relatively well-preserved fossil history, the Staphylinidae make _ interesting subjects for scientific study and they are becoming more widely used as bioindicators of environmental conditions in applied sciences such as forest research and conservation (Klimaszewski et al., 2018). A few species are even of medical importance (e.g., some Paederus spp.) or have been proposed or used in biological control (e.g., Aleochara spp.) (Thayer, 2016). This study adds to our knowledge of the distribution and life histories of the Staphylinidae of the Mid-Atlantic area of North America. NO. 53, 2019 STUDY SITES The study site includes lands managed by the National Park Service as units of the George Washington Memorial Parkway (GWMP) in Virginia (Fairfax and Arlington counties). Park sites that received the greatest inventory effort included: in Fairfax County, Dyke Marsh Wildlife Preserve, Fort Marcy, Great Falls Park, Little Hunting Creek, and Turkey Run Park and in Arlington County, Arlington Woods (at Arlington House), Gravelly Point Park, and Gulf Branch. A map of these sites is provided in Steury (2011). This area covers approximately 1,615 ha. Great Falls and Turkey Run parks and Fort Marcy and Gulf Branch fall within the Piedmont physiographic province while all other collection sites are on the Coastal Plain. Most sites are situated along the shore of the Potomac River, and the Piedmont sites border the Potomac River Gorge, an area known for high species richness of plants and animals (Cohn, 2004). Most of the study sites are dominated by maturing, second growth, primarily upland, deciduous woodlands. However, older-age stands, with dominant trees over 100 years old, occur on ridges at both the northern and southern ends of Great Falls Park. Abrams & Copenheaver (1999) documented White Oak (Quercus alba L.) individuals between 208 and 251 years old and a Black Gum (Nyssa sylvatica Marshall) 166 years old along the northern ridge. Counts of radial growth rings in 1994 on a Shortleaf Pine (Pinus echinata Mill.) that fell from a ridge along the southeastern edge of the Park dated to at least 220 years old (R. Simmons pers. comm. 2007). More open habitats can be found in moist, narrow, herbaceous dominated bands along the shore of Potomac River, in the freshwater, tidal, swamp and marsh habitats at Dyke Marsh, and in areas with managed turf grass and scattered large trees, such as Collingwood Picnic Area and Gravelly Point Park. The vascular flora of the GWMP is diverse, with more than 1,313 taxa recorded, 1,020 from Great Falls Park alone (Steury et al., 2008; Steury, 2011). MATERIALS AND METHODS Specimens were collected during a 21-year period (1998-2018) using a variety of sporadic survey methods targeting arthropods, including: Malaise traps, Lindgren funnels, blacklight (UV) bucket traps, blacklight shone on sheets, leaf litter samples processed in Berlese funnels, pitfall traps, and hand picking (including splashing along river shoreline). Six Townes. style Malaise traps (Townes, 1962) were set at Dyke Marsh, April 1998-December 1999, three each at Great Falls and Turkey Run parks (March 2006-December 2009), and BRATTAIN ET AL.: ROVE BEETLES 29: four at Little Hunting Creek (March-November 2017 and 2018). Traps at Dyke Marsh were set each year in the same locations in open, tidal, freshwater marsh dominated by Typha angustifolia L.; floodplain forest dominated by Red and Silver maple (Acer rubrum L. and A. saccharinum L.) and Tulip Poplar (Liriodendron tulipifera L.); and at the marsh/forest ecotone. In Great Falls Park, a trap was set in each of three habitats: quarry site (dry, upland, mixed deciduous/coniferous forest), swamp (dominated by Red Maple), and floodplain forest (dominated by oaks [Quercus sp.], and Tulip Poplar). In Turkey Run Park, one trap was set in upland forest dominated by oaks and tulip poplar and two traps in floodplain forest along the Potomac River (dominated by oaks, Basswood [Tilia americana L.|, and Sycamore [Platanus occidentalis L.]). At Little Hunting Creek, four traps were set in upland forest dominated by an ericaceous understory and a canopy of oaks, hickory (Carya sp.), American Beech (Fagus grandifolia Ehrh.), and some Virginia Pine (Pinus virginiana Mill.). Additional collections were also made _ by sporadically using other collecting methods, including running pitfall traps set at Dyke Marsh (five years), at Little Hunting Creek, and Great Falls and Turkey Run parks (three years), and Arlington Woods and Gulf Branch (two years); Lindgren funnel and blacklight (UV) bucket traps set at Dyke Marsh, Great Falls Park, Little Hunting Creek, and Turkey Run Park (two years); blacklight shone on sheets at Great Falls and Turkey Run parks (three years); leaf litter from Arlington Woods, Dyke Marsh, Fort Marcy, Gravelly Point Park, Great Falls Park, Gulf Branch and Turkey Run Park, processed in Berlese funnels (two years) and collecting by hand at all sites, over seven years. Hand collecting was aided by splashing (pouring water over shoreline and gravel bar habitats) unless otherwise indicated in the list of species. Locations, habitat descriptions, and collection methods are summarized in Table 1. Individuals who collected specimens of staphylinid beetles from GWMP included C. Acosta, E. Barrows, J. Brown, C. Davis, A. Evans, 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 (hp), often aided by splashing water along river shoreline. Latitude/ Eile tude Habitats sr — eer or Collection Types Arlington | Arlington County sd Arlington Woods (AW) ae o ———— oo funnels (bf), = | ae Upland, deciduous forest —— iapstpe Gravelly Point Park (GP) st oe Turf grass and river shoreline | Berlese funnels funnels 38.924 Upland, deciduous forest Gulf Branch —_—— ___ ae 115 Berlese funnels, —e'omM0 - traps | Fairfax County Count Dyke Marsh Wildlife Preserve (DM) Fort Marcy (FM) =e is Upland, deciduous forest Upland, dry, mixed deciduous/coniferous forest; deciduous swamp; deciduous floodplain forest Great Falls Park (GF) Little Hunting Creek (LH) Turkey Run Park (TR) Tidal, freshwater marsh; floodplain forest; marsh/forest Upland deciduous forest with some pine Upland deciduous forest; floodplain deciduous forest Ee funnels; black-light (UV) bucket traps (bt); Lindgren funnels (If); pitfall traps; Townes ecotone . style Malaise traps (mt) Berlese funnels Berlese funnels; black-light shone on sheets (bl); black-light (UV) bucket traps; Lindgren funnels; pitfall traps; Townes style Malaise traps Black-light (UV) bucket traps; Lindgren funnels; pitfall traps; Townes style Malaise traps Berlese funnels; black-light shone on sheets; black-light (UV) bucket traps; Lindgren funnels; pitfall traps; Townes style Malaise traps 30 BANISTERIA J. Fisher, S. Lingafelter, D. Mead, E. Oberg, M. Skvarla, D. Smith, W. Steiner, B. Steury, J. Swearingen, and C. Wirth. Specimens were determined by R. M. Brattain, A. J. Brunke, D. S. Chandler, A. V. Evans, M. L. Ferro, C. Francois, C. W. Harden, E. R. Hoebeke, J. E. Louderman, A. Marsh, A. F. Newton, B. W. Steury, and M. K. Thayer. Identifications of taxa outside the specialties of the various determiners were made with the assistance of general identification guides to all or part of North American Staphylinidae, including Downie & Arnett (1996), Newton et al. (2000), Klimaszewski et al. (2018), and Smetana (1995), supplemented by generic revisions such as Campbell (1976, 1982) and many other revisions or published notes, most of which are listed under the relevant genus in Newton et al. (2000). In addition, when possible, identifications were made or confirmed by comparison of GWMP specimens with specialist-identified specimens in the Field Museum of Natural History. In spite of these considerable efforts, the current state of knowledge of staphylinid taxonomy in North America, where many subfamilies and genera are still in need of modern revision and/or lack specialists who can identify them, has precluded the complete identification of the GWMP specimens, with the result that some species could be identified reliably only to genus (indicated as [Genus] “sp.” in the list below). New state record determinations are based on examination of the collections and the literature given in the Appendix. Specimens from GWMP were pinned, labeled, and deposited in the collections maintained at the GWMP, Turkey Run Park Headquarters in McLean, Virginia. Determiners and collection depositories for new state records are given in the Appendix. RESULTS AND DISCUSSION Two-hundred and nine taxa (171 identified to species level), in 111 genera, of staphylinid beetles were documented from GWMP. The most species rich subfamilies were Staphylininae (47 taxa), Tachyporinae (33), and Aleocharinae (29). The tribes with the most taxa were Staphylinini (43), Tachyporini (20), and Mycetoporini (13). The most species-rich genera were Philonthus (14), Sepedophilus (10) and Lordithon, Platydracus, and Stenus (7 each). The most abundant species collected in the survey area were Philonthus asper (51), Platydracus maculosus (36), Sepedophilus crassus (33), Cyparium concolor (30), Tachinus fimbriatus (27), Bryoporus — rufescens (26), Achenomorphus corticinus (24) and Oxybleptes kiteleyi (23). In order to determine which species documented from GWMP are new to Virginia a checklist was created based on literature reviews and searches of multiple NO. 53, 2019 North American collections (see appendix) documenting 558 species from Virginia. Based on the appendix, 52 species, plus five additional genera, documented from the GWMP are first records for Virginia. Additionally, 62 species new to Virginia were documented in other collections (see appendix), rendering a total of 114 species and 34 genera new to Virginia. Range extensions are documented for 16 species (see list of species for details). Of the 167 taxa documented from Piedmont sites along the Potomac River Gorge at Great Falls or Turkey Run parks or Fort Marcy or Gulf Branch during this study, 164 are first records for the gorge (Brown, 2008). Cyparium flavipes LeConte and Scaphidium obliteratum LeConte reported from the Potomac Gorge by Brown (2008) have been synonymized with other species documented from the gorge. The GWMP sites with the highest species richness were Turkey Run Park with 125 taxa, Great Falls Park (106), and Dyke Marsh Wildlife Preserve (66). Thirteen species found in GWMP are considered adventive (non-native) to Virginia and 60 species are adventive to the area comprising Virginia, Maryland, and the District of Columbia (see Appendix). Malaise traps proved to be the most successful method of capturing staphylinid beetles during this study, yielding 150 taxa. Comparable figures for other sampling methods were: hand picking (including the use of splashing), 69; Berlese funnels, 40; and pitfall traps, 24. Despite 21 years of sporadic survey effort using seven collecting techniques, 78 taxa (37.3%) documented by this study are represented by a single specimen, indicating that the list of GWMP staphylinids is very preliminary and much remains to be learned concerning the fauna of the parkway and of Virginia. LIST OF SPECIES Genera and species are listed alphabetically by subfamily, and then by supertribe, tribe and subtribe (when these are used within a subfamily). An exclamation mark is used to mark taxa not previously documented in Virginia. Adventive (non-native) species are marked with a dagger (1). The number of specimens in the collection at GWMP is indicated in parentheses after each taxon. Collection sites and methods are given using the abbreviations listed in Table 1. Other locations or collection methods are spelled out if necessary. The periods of adult activity are based on dates when live individuals were captured in the park. Dates separated by a hyphen indicate that the taxon was documented on at least one day during each month within this continuum of months, whereas dates separated by a comma represent individual observation dates. For traps set over multiple weeks, the first day of the set 1s used as the earliest date and the last day of the set as the latest date. BRATTAIN ET AL.: LIST OF SPECIES Family Staphylinidae Subfamily Aleocharinae Tribe Aleocharini Subtribe Aleocharina + Aleochara lata Gravenhorst — (4); DM, GF, TR; 11 Apr-8 Aug; mt, pf. Aleochara lustrica Say — (6); DM, GF, TR; 3 Jul-4 Sep; mt. ! Aleochara ocularis Klimaszewski — (2); GF; 10-30 Apr; mt. This record documents a southeastern range extension from Pennsylvania and Kentucky. ! Aleochara rubripes Blatchley — (1); TR; 1-20 May; mt. This specimen represents a southeastern range extension from Pennsylvania and Kentucky. ! Aleochara verna Say — (2); DM, GF; 13-20 Jun; bee bowl, mt. Fig. 1. Tribe Athetini Subtribe Athetina Atheta sp. — (3); DM, LH, TR; 26 May-7 Jul; hp, mt, pf. | Strigota cf. ambigua (Erichson) — (1); LH; 10 Jun; hp. This is the first record of this genus from Virginia, Maryland, or the District of Columbia. Subtribe Athetini incertae sedis ! Apalonia seticornis Casey — (1); AW; 14 May; bf. This record represents a northeastern range extension from Florida and Kansas. Tribe Falagriini Aleodorus bilobatus Say — (1); TR; 5 Sep-21 Oct; mt. Borboropora quadriceps (LeConte, J. L.) — (1); TR; 7- 21 Jun; mt. Myrmecocephalus cingulatus (LeConte, J. L.)—(2); DM, TR; 6-20 Jun, 18 Aug-4 Sep; mt. Myrmecocephalus concinnus (Erichson) — (1); DM; 20 Jun-1 Jul; mt. Tribe Geostibini Aloconota sp. — (3), GF, TR; 21 May-17 Jul; mt. Geostiba sp. — (1); FM; 15 Apr; bf. ROVE BEETLES 31 Tribe Homalotini Subtribe Bolitocharina Leptusa sp. — (2); GF, TR; 5 Sep-1 Dec; mt. ! Phymatura cf. blanchardi (Casey) — (4); GF, TR; 9 Sep-1 Dec; mt. This is the first record of this genus recorded from Virginia, Maryland, or the District of Columbia. This Phymatura species is difficult to identify but is most likely blanchardi when compared to previously named museum specimens and referenced to available literature. Subtribe Gyrophaenina Gyrophaena sp. — (3); DM, GF; 18 May-30 Jul; hp, mt. Phanerota fasciata (Say) — (2); GF, TR; 19 Jun-17 Jul; mt Tribe Hoplandriini Subtribe Hoplandriina Hoplandria laevicollis (Notman) — (3); TR; 19-30 Jun; mt. Hoplandria sp. — (6); GF, TR; 21 May-30 Jul; mt, pf. Tribe Hypocyphtini Oligota cf. pusillima (Gravenhorst) — (1); GP; 14 May; bf. If identified correctly, this species is believed to be adventive in North America, known from Canada (New Brunswick) and the United States (Massachusetts and New York). Tribe Lomechusini Subtribe Myrmedoniina Pella sp. — (3); DM; 15 Apr-18 May; hp. Tribe Myllaenini Myllaena sp. — (2); DM, TR; 20 Jun-4 Sep; mt. Tribe Oxypodini Subtribe Oxypodina ! Gennadota sp. — (3); TR; 9 Sep-17 Nov; mt. This is the first record of this genus from Virginia, Maryland, or the District of Columbia. 32 BANISTERIA Oxypoda cf. opaca (Gravenhorst) — (3); DM, TR; 21 Jul- 4 Aug, 21 Nov-31 Jan; mt. Even when compared to previously named museum specimens and referenced to available literature, this species is difficult to identify but is most closely allied to O. opaca. Oxypoda sp. — (2); AW, DM; 14-18 May; bf, hp. ! Tetralaucopora sp. — (1); DM; 7-21 Nov; mt. This is the first record of this genus from Virginia, Maryland, or the District of Columbia. Tribe Placusini Placusa sp. — (7); DM, TR; 6 Jun-4 Sep; mt. Tribe Tachyusini Gnypeta cf. nigrella (LeConte, J. L.)— (12); TR; 14 Jun; hp. This species is difficult to identify without male genitalia, but is probably G. nigre/la when compared to previously named museum specimens and referenced to available literature. Tachyusa sp. — (2); GF, TR; 4-18 May; hp. Subfamily Euaesthetinae Tribe Euaesthetini Euaesthetus cf. americanus Erichson — (3); DM; 15 Apr- 18 May; hp. This Euaesthetus is difficult to identify but is most closely allied to E. americanus when compared to previously named museum specimens. Euaesthetus iripennis Casey — (1); AW, 15 Apr, bf. Subfamily Megalopsidiinae Megalopinus caelatus (Gravenhorst) — (2); GF, LH; 2-30 Jun; mt. Megalopinus rufipes (Motschulsky) — (2); GF, LH; 13 Apr-15 May; mt. Subfamily Omaliinae Tribe Anthophagini Acidota subcarinata Erichson — (1); TR; 22 Oct-1 Dec; mt. Arpedium schwarzi Fauvel — (7); DM, GF, TR; 15 Apr- 7 Jul, 22 Oct-1 Dec; hp, mt, pf. Arpedium n. sp. — (1); DM; 24 Oct-8 Nov; mt. Fig. 2. Recognized as undescribed by Margaret Thayer based on male genitalia, this new species is also known from North Carolina, Ohio, and Oklahoma. It is being described by Alexey Shavrin in a revision of North American species of Arpedium. NO. 53, 2019 Geodromicus brunneus (Say) — (13); GF, LH, TR; 24 Apr-25 May, 22 Oct-1 Dec; bl, hp, mt. Lesteva pallipes LeConte, J. L. — (6); DM, GF, TR; 27 Apr-20 May, 22 Oct-1 Dec; If, mt. Olophrum obtectum Erichson — (6); AW, DM, TR; 18 Mar-14 May, 21 Nov-5 Dec; bf, mt. Trigonodemus striatus LeConte, J. L. — (3); GF, TR; 5 Sep-17 Nov; mt. This species was first reported from Virginia by Steury (2017). Tribe Omaliini Hapalaraea hamata (Fauvel) — (1); TR; 5 Sep-21 Oct; mt. ! Omalium (sensu lato) fractum Fauvel — (1); TR; 22 Oct- 1 Dec; mt. Fig. 3. (generic placement: see Newton et al., 2000: 338, under Pycnoglypta). This first Virginia record fills a gap in the broad distribution extending northwest to Illinois and Michigan; south-southwest to Texas and Oklahoma, Georgia, Kentucky, and North Carolina and northeast to New York. ! Omalium repandum Erichson — (1); TR; 22 Oct-17 Nov; mt. This specimen fills a range gap in a broad distribution extending northeast, west, and northwest. ! Phyllodrepa humerosa (Fauvel) — (2); DM, GF; 27 Oct- 5 Dec; mt. These Virginia records fill a gap in the broadly documented distribution of this species which extends west to Wisconsin through Oklahoma, south to Georgia, and north to Pennsylvania and Nova Scotia, Canada. Phyllodrepa punctiventris (Fauvel) — (2); AW; 14 May; bf. ! + Xylodromus concinnus (Marsham) — (1); GF; Apr 11- 28; If. This first Virginia record adds to a poorly documented distribution scattered over North America. Subfamily Osoriinae Tribe Eleusinini ! Eleusis pallida (LeConte, J. L.) — (1); TR; 1-20 May; mt. Tribe Osoriini Molosoma latipes (Gravenhorst) — (1); TR; 13 Jun; hp. Tribe Thoracophorini Subtribe Clavilispinina ! Clavilispinus rufescens (LeConte, J. L.)—(2); AW, LH; 13 Apr-15 May; bf, mt. Fig. 4. These records document a northern range extension from South Carolina. BRATTAIN ET AL.: Subtribe Thoracophorina Thoracophorus costalis (Erichson) — (2); AW; 15 Apr- 14 May; bf. Subfamily Oxyporinae Oxyporus stygicus Say — (1); TR; 1-20 May; mt. Pseudoxyporus lateralis (Gravenhorst) — (1); TR; 21 Jul- 4 Aug; mt. Pseudoxyporus occipitalis (Fauvel)— (1); TR; 1-20 May; mt. Pseudoxyporus quinquemaculatus (LeConte, J.L.) — (17); GF; 21 May-30 Jun; mt. Twelve of these were captured in the same Malaise trap sample set in Great Falls Swamp. Subfamily Oxytelinae Tribe Blediini Bledius cf. annularis LeConte, J. L. — (1); TR; 4 May; hp. More than 28 species of this genus are reported from the Northeast. Some of the distinguishing characters are subtle in structure, making the species difficult to identify. Bledius semiferrugineus LeConte, J. L. — (2); TR; 3-4 May; hp. Tribe Oxytelini +t Anotylus rugosus (Fabricius) — (3); DM, GF; 11 Apr- 23 May; mt, pf. Anotylus sp. — (4); AW, TR; 15 Apr, 30 Jun-24 Sep; bf, pf. ! Apocellus sphaericollis (Say) — (9); AW, Collingwood Picnic Area, GF, TR; bf, hp (along curb of parking lot), mt, pf. + Oxytelus laqueatus (Marsham) — (1); TR; 31 Jul-17 Aug; mt. ! Oxytelus pensylvanicus Erichson — (1); GF; 1-15 Jul; mt. Fig. 5. ! + Oxytelus sculptus Gravenhorst — (1); TR; 18 Aug-4 Sep; mt. Tribe Thinobiini ! + Carpelimus bilineatus Stephens — (7); DM, GF, TR; 15 Apr-14 Jun; hp, mt. ! Carpelimus quadripunctatus (Say) — (5); DM, GF, TR; 1 May-1 Aug; bl, hp, mt. Carpelimus sp. — (1); DM; 15 Apr; hp. | Thinodromus arcifer (LeConte, J. L.)—(1); DM; 19-28 Apr; mt. Fig. 6. ROVE BEETLES BB Subfamily Paederinae Tribe Lathrobiini Subtribe Lathrobiina Lathrobium sp. — (1); TR; 1-20 May; mt. ! Lobrathium collare (Erichson) — (3); TR; 25 May, 9 Sep-21 Oct; hp, mt. Pseudolathra sp. — (2); DM, LH; 10 Jun-8 Aug; hp, mt. Tetartopeus sp. — (1); GF; 5 Jul; bl. Subtribe Medonina Achenomorphus corticinus (Gravenhorst) — (24); AW, DM, GF, TR; 15 Apr- 30 Oct; bf, bt, mt. ! Sunius confluentus (Say) — (5); GF, LH; 13 Apr-15 May; hp (on tree trunk at night, under bark), mt. | Sciocharis carolinensis Casey — (1); GF; 19-30 Jun; mt. Fig. 7. | Sciocharis exilis (Erichson) — (1); AW; 15 Apr; bf. Fig. 8. This specimen documents a northern range extension from Alabama and Florida. Subtribe Scopaeina Scopaeus sp. —(8); DM, GF, TR; 15 Apr-18 May, 5 Aug- 22 Oct; hp, mt. Subtribe Stilicina Eustilicus tristis (Melsheimer, F. E.) — (3); TR; 1 May-5 Jun; mt. !Rugilus angularis (Erichson) — (1); GF; 15 Apr; bf. Subtribe Stilicopsina Stamnoderus monstrosus (LeConte, J. L.) — (1); DM; 18 May; hp. Tribe Paederini Subtribe Cryptobiina Homaeotarsus badius (Gravenhorst) — (13); GF, TR; 24 Apr-7 Sep; hp, mt. Homaeotarsus bicolor (Gravenhorst) — (8); GF, TR; 20 May-17 Jul, 22 Oct-1 Dec; bt, hp, mt. Homaeotarsus capito (Casey) — (2); AW, GF; 11-28 Apr; bf, If. Homaeotarsus carolinus (Erichson) — (1); GF; 11-28 Apr; pf. ! Homaeotarsus cribratus (LeConte, J. L.) — (2); TR; 19- 21 Sep; pf. NO. 53, 2019 BANISTERIA 34 Figs. 1-4 (left to right). Figs. 5-8 (left to right). BRATTAIN ET AL.: ROVE BEETLES 35 ! Homaeotarsus pimerianus (LeConte, J. L.) — (2); GF; 31 July; hp. Figs. 9-10. These specimens represent a significant eastern range extension from Indiana and Texas. Subtribe Paederina Paederus obliteratus (LeConte, J. L.) —(1); DM; 15 Apr; bf. Tribe Pinophilini Subtribe Pinophilina Pinophilus latipes Gravenhorst — (2); GF; 23 Jun; bl. Subfamily Piestinae | Siagonium americanum (Melsheimer) — (1); LH; 3-16 Jun; If. This first Virginia record fills a gap in a broad eastern distribution in North America. Subfamily Proteininae Tribe Proteinini Proteinus sp. — (19); LH; 14-30 Jun; mt. This species is similar to the adventive species P. atomarius Erichson, reported from the District of Columbia, but the identity Figure captions (opposite page) could not be confirmed based on the single female. Subfamily Pselaphinae Supertribe Batrisitae Tribe Batrisini Subtribe Batrisina Arthmius globicollis LeConte, J. L. — (5); DM, GF, TR: 14 Mar-15 Apr, 5 Aug-21 Oct; bf, hp, mt. ! Batrisodes furcatus (Brendel) — (2); TR; 23 May-5 Jun; mt. Fig. 11. This species has been documented north to New York and south to Alabama, so it was to be expected in Virginia. Batrisodes lineaticollis (Aubé) — (1); GF; 21 May-18 Jun; mt. ! Batrisodes scabriceps (LeConte, J. L.) — (1); TR; 21 Jun-6 Jul; mt. Batrisodes schaumii (Aubé) — (1); TR; 15-30 Oct; mt. ! Batrisodes striatus (LeConte, J. L.) — (1); GF; 11-28 Apr; pf. Supertribe Euplectitae Tribe Euplectini Euplectus confluens LeConte, J. L. — (3); AW, FM; 15 Apr-14 May; bf. Fig. 1. Aleochara verna Say — Dyke Marsh Wildlife Preserve, 13-20 June 1999, Malaise trap. Collected by E. Barrows. Determined by E. R. Hoebeke, Length 4.3 mm. Fig. 2. Arpedium new species — Dyke Marsh Wildlife Preserve, 24 October-8 November 1999, Malaise trap. Collected by E. Barrows. Determined by M. K. Thayer. Length 4.1 mm. Fig. 3. Omalium (sensu lato) fractum (Fauvel) — Turkey Run Park, 22 October-1 December 2009, Malaise trap. Collectors D. Smith & B. Steury. Determined by M. K. Thayer. Length 2.2 mm. Fig. 4. Clavilispinus rufescens (LeConte, J. L.) — Little Hunting Creek, 13 April-15 May 2017, Malaise trap. Collectors C. Acosta, C. Davis, & B. W. Steury. Determined by R. M. Brattain. Length 3.5 mm. Fig. 5. Oxytelus pensylvanicus Erichson — Great Falls Park, quarry site, 1-15 July 2008, Malaise trap. Collector D. Smith. Determined by R. M. Brattain. Length 2.4 mm. Fig. 6. Thinodromus arcifer (LeConte, J. L.)— Dyke Marsh Wildlife Preserve, 19-28 April 1998, Malaise trap. Collector E. Barrows. Determined by R. M. Brattain. Length 2.5 mm. Fig. 7. Sciocharis carolinensis Casey — Great Falls Park quarry site, 19-30 June 2009, Malaise trap. Collectors D. Smith & B. Steury. Determined by R. M. Brattain. Length 3.5 mm. Fig. 8. Sciocharis exilis (Erichson) — Arlington Woods, Berlese funnel, 15 April 2013. Collectors M. S. Skvarla & J. R. Fisher. Determined R. M. Brattain. Length 2.5 mm. 36 BANISTERIA NO. 53, 2019 Figs. 9-12 (left to right). Figs. 9-10. Homaeotarsus pimerianus (LeConte, J. L.) — Great Falls Park at mouth of Difficult Run along Potomac River, hand collected by splashing water on silty gravel shoreline, 31 July 2017. Collector. B. W. Steury. Determined by R. M. Brattain & A. F. Newton. Length 9.8 mm. Left, anterior dorsal habitus, right, close up of head and pronotum. Fig. 11. Batrisodes furcatus (Brendel) — Turkey Run Park, Malaise trap, 23 May-6 June 2008. Collector D. Smith. Determined by D. S. Chandler. Length 2.1 mm. Fig. 12. Neobisnius paederoides (LeConte, J. L.) — Turkey Run Park, hand collected by splashing water on silt-caked leaf-litter along shore of Potomac River, | May 2017. Collector B. W. Steury. Determined by A. Brunke. Length 5.0 mm. Tribe Trichonychini Subtribe Panaphantina Eutyphlus similis LeConte, J. L. — (1); GF; 14 Mar; hp. Subtribe Trimiina Melba parvula (LeConte, J. L.) — (2); DM; 18 May; A iue Ms dubia (LeConte, J. L.) — (2); 16 May; FM; bf. Tribe Trogastrini Subtribe Rhexiina Rhexius schmitti Brendel — (3); AW, FM, LH; 13 Apr-16 May; bf, mt. Rhexius substriatus LeConte, J. L. — (2); DM. TR; 1-20 May; hp, mt. Supertribe Goniaceritae Tribe Brachyglutini Subtribe Brachyglutina Brachygluta abdominalis (Aubé) — (2); GF, TR; 18 Aug- 30 Oct; mt. Brachygluta luniger (LeConte, J. L.) — (7); DM, GF; 15 May-29 Jun; bf, mt. Brachygluta ulkei (Brendel) — (12); DM; 15 Apr-18 May; bf. Subtribe Pselaptina Eutrichites zonatus (Brendel) — (1); DM; 15 Apr; bf. BRATTAIN ET AL.: ROVE BEETLES 37 Supertribe Pselaphitae Tribe Ctenistini Ctenisodes sp. — (1); DM; 15 Apr; bf. Tribe Tmesiphorini Tmesiphorus carinatus (Say) — (1); DM; 6 Jun; bf. Tribe Tyrini Subtribe Tyrina ! Cedius spinosus LeConte, J. L. — (1); LH; 19 May-2 Jun; mt Subfamily Scaphidiinae Tribe Cypariini ! Cyparium concolor (Fabricius) — (30); DM, GF, TR; 21 May-7 Sep; mt. Tribe Scaphidiini Scaphidium piceum Melsheimer, F. E. — (13); GF, TR; 21 May-30 Jul; hp, mt, | Scaphidium quadriguttatum Say — (3); GF, LH, TR; 21 May-18 Jun, 11-28 Aug; hp (under log), mt. Tribe Scaphisomatini Baeocera sp. 1 — (2); TR; 1 May-30 Jun; mt. Baeocera sp. 2 — (3); DM, GF; 16 Jul-7 Sep; mt. Scaphisoma convexum Say — (3); GF, TR; 21 May-30 Jul; mt. ! Scaphisoma terminatum Melsheimer, F. E. — (1); GF; 31 Jul-17 Aug; mt. Scaphisoma sp. | — (2); GF; 30 Jun-25 Aug; mt. Scaphisoma sp. 2 — (1); TR; 3-17 Jul; mt. Toxidium gammaroides LeConte, J. L. — (2); GF, TR; 16 Jul-17 Aug; mt. Subfamily Scydmaeninae Supertribe Cephenniitae Tribe Cephenniini Cephennodes cf. virginicus (Casey) — (6); AW, FM, GB; 15 Apr, 19-21 Oct; bf. Based on known ranges of closely allied species this taxon is most likely C. virginicus but it could be an undescribed species. Supertribe Scydmaenitae Tribe Chevrolatiini Chevrolatia amoena LeConte, J. L — (1); LH; 1-14 Jun; mt. Tribe Glandulariini Euconnus spp. — (8); AW, GF, Riverside Park, TR; 13 Apr-6 Jul, 19 Sep-21 Oct; bf, hp, mt. ! Parascydmus sp. — (1); LH; 19 Sep-10 Oct; mt. This genus has not been previously recorded from Virginia, Maryland, or the District of Columbia. Subfamily Staphylininae Tribe Diochini Diochus schaumii Kraatz — (4); DM, GF; 15 Apr. bf, hp. Tribe Staphylinini Subtribe Acylophorina ! Acylophorus agilis Smetana — (4); DM; 19 Apr-9 Aug; mt. ! Acylophorus caseyi Leng — (2); GF; 1 May, 15 Aug-7 Sep; hp, mt. These first Virginia records represent a range extension for this mainly northern species which has been known from West Virginia and Pennsylvania to the north but has also been documented south to Alabama and Louisiana. Acylophorus flavicollis Sachse — (1); GF; 1 Jun; hp (edge of vernal pool). | Hemiquedius infinitus Brunke & Smetana — (1); GF; 5 Sep-21 Oct; mt. Subtribe Anisolinina Tympanophorus puncticollis (Erichson) — (1); GF; 19-30 Jun; mt. Subtribe Erichsoniina Erichsonius alumnus Frank, J. H. — (9); DM, GF, TR; 12 Apr-11Jun; hp (swamp), mt. Erichsonius brachycephalus Frank, J. H. — (4); GF, TR; 21 May-17 Jul; mt. Subtribe Philonthina Belonuchus rufipennis (Fabricius) — (19); DM, GF, TR; 14 Apr-26 Sep; hp, mt, pf. 38 BANISTERIA Bisnius blandus (Gravenhorst) — (5); AW, GF, LH, TR; 15 Apr-18 May, 3 Jul-4 Sep; bf, mt, pf. | Bisnius pugetensis (Hatch) —(1); TR; 21 Jul-4 Aug; mt. This first Virginia record represents a southern range extension from Pennsylvania. Hesperus apicialis (Say) — (9); GF, LH, TR; 10 Apr-11 Aug; hp, If, mt. Hesperus baltimorensis (Gravenhorst) — (12); DM, GF, TR; 11 Apr-18 May, 1 Jul-26 Oct; hp, lf mt. ! Hesperus stehri Moore — (1); TR; 28 Apr-12 May; pf. This specimen represents a northeastern range extension from North Carolina and Tennessee. Laetulonthus laetulus (Say) — (2), GF; 11 Apr-19 May; If. Neobisnius jocosus (Horn) — (6); DM, GF, TR; 21 May- 4 Sep; mt. ! Neobisnius paederoides (LeConte, J. L.) — (4); TR; 1-4 May; hp. Fig. 12. Neobisnius sobrinus (Erichson) — (3); DM, TR; 28 Apr- 10 May, hp, mt. Neobisnius terminalis (LeConte, J. L.) — (1); TR; 4 May, hp. Philonthus asper Horn — (51); AW, DM, GF, TR; 15 Apr-13 Oct; bf, hp, If, mt, pf. Philonthus caeruleipennis (Mannerheim) — (6); GF, TR; 21 May-21 Jul, 26 Sep; hp. mt. + Philonthus cognatus Stephens — (2); TR; 1-21 Jul; mt. + Philonthus debilis (Gravenhorst) — (1); TR; 19-21 Sep; pf. Philonthus gracilior Casey — (2); GF, TR; 11-13 Jun; hp (swamp and splashing). Philonthus lomatus Erichson — (7); DM, GF, TR; 28 May-21 Jul, 26 Sep; hp, mt. ! Philonthus palliatus (Gravenhorst) — (1); DM; 6-20 Jul; mt. Philonthus quadricollis Horn — (2); TR; 1 May; hp. Philonthus rufulus Horn — (21); DM, GF, LH, TR; 10 Apr-21 Jul, 15-30 Oct; bl, hp (under log and splashing), mt, pf. Philonthus sericans (Gravenhorst) — (21); DM, GF, TR; 19 Apr-21 Oct; hp, mt. Philonthus thoracicus (Gravenhorst) — (1); LH, 10 Jun; hp. ! Philonthus umbrinoides Smetana — (1); DM; 8-23 May; mt. This specimen documents a southern range extension from New York and West Virginia. Philonthus validus Casey — (6); DM, GF, TR; 7 Jul-21 Oct; mt. ! + Philonthus ventralis (Gravenhorst) — (1); TR; 18 Mar- 9 Apr; mt. NO. 53, 2019 Subtribe Quediina Quedius capucinus (Gravenhorst) — (3); GF; 19 Sep-21 Oct; mt. + Quedius mesomelinus (Marsham) — (1); TR; 16-30 Jul; mt. Quedius peregrinus (Gravenhorst) — (3); GF, TR; 21 May-21 Jul, 19 Sep-21 Oct; mt. Subtribe Staphylinina Platydracus cinnamopterus (Gravenhorst) — (8); AW, FM, GF, TR; 14 Apr-26 Jul; bf, hp (under bark), mt. ! Platydracus exulans (Erichson) — (1); GF; 10-30 Apr; mt. Platydracus maculosus (Gravenhorst) — (36); AW, GF, LH, TR; Feb 5-12, 1 Jun-21 Oct; hp, If, mt, pf. Platydracus praetermissus Newton — (2); LH, GB; 15 Apr, 3-16 Jun; bf, pf. ! Platydracus violaceus (Gravenhorst) — (6); LH, GF, TR; 14 Apr-15 Jul; hp (under bark), mt, pf. ! Platydracus viridanus (Horn) — (1); TR; 21 Jul-4 Aug; mt. Platydracus zonatus (Gravenhorst) — (6); AW, DM, FM, GB, TR; 15 Apr-25 May; bf, hp, pf. ! + Tasgius winkleri (Bernhauer) — (1); TR; 16 Sep; hp. This first Virginia record represents a southern range extension from Pennsylvania for this adventive species, which is slowly expanding its range in North America. Tribe Xantholinini Neohypnus emmesus (Gravenhorst) — (4); AW, DM, LH, GF; 14-29 Apr: bf, hp, If, mt. Neohypnus sp. — (2); AW, DM; 14 May, 9 Aug; bf, mt. ! Oxybleptes kiteleyi Smetana — (23); GF, TR; 16 Jul-21 Oct; mt. These first Virginia records represent a range extension for this rare species which was previously known only from New York and farther north, except for a record from North Carolina. Subfamily Steninae Stenus callosus Erichson — (10); DM, GF, TR; 1 May-4 Sep, 21 Nov-5 Dec; hp, mt. Stenus colon Say — (3); GF, TR; 24 Apr-1 Jun; hp. Stenus colonus Erichson — (6); DM, TR; 4 May-14 Jun; hp, mt. ! Stenus croceatus (Casey) — (2); DM; 18-30 May; hp, mt. Stenus egenus Erichson — (1); TR; 20 Apr; hp Stenus femoratus Say — (4); DM, TR; 1 May, 26 Sep-11 Oct; hp, mt. Stenus sp. — (1); DM; 18 May; hp. BRATTAIN ET AL Subfamily Tachyporinae Tribe Mycetoporini | + Bolitobius cingulatus Mannerheim — (1); GF; 18 Aug- 4 Sep; mt. This Virginia specimen represents a range extension for this adventive species which is also known from New Jersey and farther north, and from Alabama and Florida to the south. ! Bryophacis smetanai Campbell — (1); TR; 19-30 Jun; mt. This record represents a southern range extension from Pennsylvania. Bryoporus rufescens LeConte, J. L. — (26); DM, GF, TR; 1 May-21 Oct; mt. Bryoporus testaceus LeConte, J. L. — (1); GF; 20 May; hp. ! Lordithon appalachianus Campbell — (14); GF, TR; 1 May-1 Dec; mt. Lordithon cinctus (Gravenhorst) — (11); AW, GF, LH, TR; 15 Apr-21 Oct; bf, mt. Lordithon facilis (Casey) — (8); AW, GF, TR; 14 May- 15 Jul, 5 Sep-21 Oct; bf, mt. Lordithon kelleyi (Malkin) — (8); DM, GF, TR; 1 May- 21 Oct; mt. Lordithon niger (Gravenhorst) — (1); TR; 1-15 Jul; mt. Lordithon notabilis Campbell — (3); GF, TR; 19 Jun-21 Oct; mt. Lordithon quaesitor (Horn) — (2); TR; 19-30 Jul; mt. Mycetoporus americanus Erichson — (4); GF, TR; 10-30 Apr, 7 Jul-4 Sep; mt. Mycetoporus lucidulus LeConte, J. L. — (3); GF, TR; 10- 30 Apr, 16 Jul-17 Aug; mt. Tribe Tachyporini Coproporus laevis LeConte, J. L. — (11); GF; 25 Jun, 19 Sep-21 Oct; bl, mt. Coproporus ventriculus (Say) — (11); LH, GF, TR; 12 Apr-7 Sep; bl, hp, If, mt. ! Nitidotachinus scrutator (Gemminger & Harold) — (2); TR; 19 Jun-15 Jul; mt. Sepedophilus basalis (Erichson) — (2); FM, GF; 16 May- 29 Jun; hp, mt. Sepedophilus cinctulus (Erichson) — (1); TR; 5-25 Aug; mt. Sepedophilus crassus (Gravenhorst) — (33); Fort Hunt Park, GF, TR; 15 Jun-1 Dec; hp, mt. Sepedophilus frosti Campbell — (4); DM, GF, TR; 24 Jun-30 Jul, 22 Oct-1 Dec; mt. + Sepedophilus littoreus (Linnaeus) — (13); AW, DM, GF, TR; 10 Apr-4 Sep; mt. ! Sepedophilus occultus (Casey) — (1); GF; 15-29 Jun; mt. .. ROVE BEETLES 39 Sepedophilus opicus (Say) — (9); AW, FM, GF, TR; 15 Apr-20 Jul; bf, hp, mt. Sepedophilus scriptus (Horn) — (1); GF; 21 May-18 Jun; mt. Sepedophilus velocipes (Casey) — (2); TR; 2 Jul-4 Sep; mt. Sepedophilus versicolor (Casey) — (13); AW, LH, GF, TR; 11 Apr-21 Oct; bf, lf mt, pf. Tachinus axillaris Erichson — (3); GF, TR; 27 Apr-19 May, 7-30 Jul; mt, pf. Tachinus canadensis Horn — (3); GF, TR; 13 Oct-1 Dec; mt, pf. Tachinus fimbriatus Gravenhorst — (27); DM, GF, TR; 21 May-7 Jul, 19 Sep-5 Dec; bl, mt, pf. Tachinus fumipennis (Say) — (6); GF, TR; 7 Jun-30 Jul, 5 Sep-21 Oct; mt, pf. Tachinus minimus Campbell — (1); TR; 19-30 Jun; mt. Tachyporus jocosus Say — (1); AW; 14 May; bf. ! Tachyporus lecontei Campbell — (1); GB; 17 Jun; hp. This specimen represents a southern range extension from Pennsylvania. ACKNOWLEDGMENTS With much gratitude, we acknowledge the assistance of our dedicated cadre of citizen science volunteers working in the GWMP bug lab. Without their meticulous sorting of Malaise trap samples, this study would not have been possible. We also heartily thank the many specialists listed in the Materials and Methods and the Appendix for their identification efforts and new state record determinations. Steve Roble, Virginia Department of Conservation & Recreation, Division of Natural Heritage, provided helpful comments and formatting advice on the manuscript. LITERATURE CITED Abrams, M. L., & C. A. Copenheaver. 1999. Temporal variation in species recruitment and dendroecology of an old-growth white oak forest in the Virginia Piedmont, USA. Forest Ecology and Management 124: 275-284. Betz, O., U. Irmler, & J. Klimaszewski (eds.). 2018. Biology of Rove Beetles (Staphylinidae): Life History, Evolution, Ecology and _ Distribution. Springer International Publishing, Cham, Switzerland. 351 pp. Brown, J. W. 2008. The invertebrate fauna of Plummers Island, Maryland. Contribution XXX _ to the Natural History of Plummers Island, Maryland. Bulletin of the Biological Society of Washington 15: 1-226. 40 BANISTERIA Brunke, A. J., & J. Buffam. 2018. A review of Nearctic rove beetles (Staphylinidae) specialized on the burrows and nests of vertebrates. Pp. 145-159 In O. Betz, U. Irmler, & J. Klimaszewski (eds.), Biology of Rove Beetles (Staphylinidae): Life History, Evolution, Ecology and Distribution. Springer International Publishing, Cham, Switzerland. Campbell, J. M. 1976. A revision of the genus Sepedophilus Gistel (Coleoptera: Staphylinidae) of America north of Mexico. Memoirs of the Entomological Society of Canada 99: 1-89. Campbell, J. M. 1982. A revision of the genus Lordithon Thomson of North and Central America (Coleoptera: Staphylinidae). Memoirs of the Entomological Society of Canada 119: 1-116. Chatzimanolis, S. 2018. A review of the fossil history of Staphylinoidea. Pp. 27-45 Jn O. Betz, U. Irmler, & J. Klimaszewski (eds.), Biology of Rove Beetles (Staphylinidae): Life History, Evolution, Ecology and Distribution. Springer International Publishing, Cham, Switzerland. Cohn, J. P. 2004. The wildest urban river: Potomac River Gorge. BioScience 54: 8-14. Downie, N. M., & R. H. Arnett, Jr. 1996. The Beetles of Northeastern North America. Vols. 1, 2. Sandhill Crane Press, Gainesville, FL. 1,721 pp. Irmler, U., J. Klimaszewski, & O. Betz. 2018. Introduction to the biology of rove beetles. Pp. 1-4 Jn O. Betz, U. Irmler, & J. Klimaszewski (eds.), Biology of Rove Beetles (Staphylinidae): Life History, Evolution, Ecology and _ Distribution. Springer International Publishing, Cham, Switzerland. Klimaszewski, J., A. J. Brunke, T. T. Work, & L. Venier. 2018. Rove beetles (Coleoptera, Staphylinidae) as bioindicators of change in boreal forests and their biological control services in agroecosystems: Canadian case studies. Pp. 161-181 Jn O. Betz, U. Irmler, & J. Klimaszewski (eds.), Biology of Rove Beetles (Staphylinidae): Life History, Evolution, Ecology and Distribution. Springer International Publishing, Cham, Switzerland. Klimaszewski, J., R. P. Webster, D. W. Langor, A. Brunke, A. Davies, C. Bourdon, M. Labrecque, A. F. Newton, J.-A. Dorval, & J. H. Frank. 2018. Aleocharine rove beetles of eastern Canada NO. 53, 2019 (Coleoptera, Staphylinidae, Aleocharinae): a glimpse of megadiversity. Springer-Verlag, Cham, Switzerland. 902 pp. Newton, A. F. 2019. StaphBase: Staphyliniformia world catalog database (version Nov. 2018): Staphylinoidea, Hydrophiloidea, Synteliidae. In Y. Roskov, G. Ower et al. (eds.). Species 2000 & ITIS Catalogue of Life. Species 2000: Naturalis, Leiden, the Netherlands. Digital resource at www.catalogueoflife.org/col. Last accessed 8 February 2019. Newton, A. F., M. K. Thayer, J. S. Ashe, & D. S. Chandler. 2000. Staphylinidae Latreille, 1802. Pp. 272— 418 In R. H. Arnett, Jr. & M. C. Thomas (eds.), American Beetles. Volume I. Archostemata, Myxophaga, Adephaga, Polyphaga: Staphyliniformia. CRC Press, Boca Raton, FL. Smetana, A. 1995. Rove beetles of the subtribe Philonthina of America north of Mexico (Coleoptera: Staphylinidae). Classification, phylogeny and taxonomic revision. Memoirs on Entomology, International 3. 946 pp. Steury, B. W. 2011. Additions to the vascular flora of the George Washington Memorial Parkway, Virginia, Maryland, and the District of Columbia. Banisteria 37: 3-20. Steury, B. W. 2017. First record of the rove beetle Trigonodemus _ striatus LeConte (Coleoptera: Staphylinidae) from Virginia and additional new park records (Coleoptera: Anthicidae, Buprestidae, Carabidae, Cerambycidae, Chrysomelidae) for the George Washington Memorial Parkway. Banisteria 48: 14-16. Steury, B. W., G. P. Fleming, & M. T. Strong. 2008. An emendation of the vascular flora of Great Falls Park, Fairfax County, Virginia. Castanea 73: 123-149. Thayer, M. K. 2016. Staphylinidae Latreille, 1802. Pp. 394-442 In R. G. Beutel & R. A. B. Leschen (eds.), Coleoptera, Beetles. Vol. 1: Morphology and Systematics (Archostemata, Adephaga, Myxophaga, Polyphaga partim). 2nd edition. Handbook of Zoology; Arthropoda: Insecta (R. G. Beutel & N. P. Kristensen, eds.). De Gruyter, Berlin/Boston, Florida. Townes, H. 1962. Design for a Malaise trap. Proceedings of the Entomological Society of Washington 64: 253-— 262. BRATTAIN ET AL.: ROVE BEETLES 4] APPENDIX Preliminary Checklist of Rove Beetles (Coleoptera: Staphylinidae) of Virginia, Maryland, and the District of Columbia Alfred F. Newton (compiler) The following checklist includes all species or subspecies of Staphylinidae that are currently known to occur in Virginia, Maryland and the District of Columbia, including those from Virginia newly reported in this work. The list contains 792 taxa including 558 from Virginia, 343 from Maryland, and 494 from the District of Columbia. Sixty species are believed to be adventive in North America, originating from the Palaearctic region and in most cases probably from Europe, with the exception of Anotylus insignitus from the Neotropical region. Another fifteen species are Holarctic (or even more broadly distributed). One hundred and fourteen species are first records for Virginia, eight are new to Maryland and four are new to the District of Columbia. The Checklist is extracted from a world catalog database compiled and maintained by AEN, which is now available online in simplified form via the Catalogue of Life (Newton 2019). The inclusion of these states and district in the distribution of these taxa is based, with rare exceptions, on published sources, which include the original descriptions of each name (all were consulted), and secondary sources including the catalogs of Horn (1868), Ulke (1902), Leng (1920), Blackwelder (1973a, 1973b), Moore & Legner (1975), Chandler (1997), Herman (2001), Gusarov (2003) and Lobl (2018), recent regional reviews including Downie & Arnett (1996), Newton et al. (2000), O’Keefe (2000), Brown (2008), Brunke et al. (2011), and Klimaszewsk1 et al. (2013, 2018), and generic revisions, monographs, notes or similar sources concerning many taxa, most of which are listed in the individual generic treatments for Staphylinidae in American Beetles (Newton et al. 2000, O’Keefe, 2000). A few previously unreported state records (marked below as new) are included based on still-unpublished revisionary work or identifications by AEN or others as indicated, and additional new records (also marked as such) are included based on the above GWMP study. In spite of efforts to make this list as complete and accurate as possible, there are undoubtedly additional published records that were overlooked, and some taxa included in the list may have been erroneously reported from these areas. Based on what is known about the overall distribution patterns of staphylinids in the eastern United States, many additional species can be expected to actually occur in these states or district but, to our knowledge, have not yet been found or reported from there; the actual staphylinid diversity of this area is no doubt much larger than shown in this list. Thus, we consider this checklist a preliminary one, in need of further review and documentation. The format of the checklist is adapted from Bousquet et al. (2013). Further information about each species or subspecies listed here, including the original generic combination (if different from the current one), a reference to the original description, synonyms (if any), and approximate overall distribution, can be found in the Catalogue of Life (Newton, 2019). Abbreviations Areas: DC District of Columbia MD Maryland (state) VA Virginia (state) Identifiers (for new state or district records): AFN Alfred F. Newton, FMNH, Chicago, Illinois AJB Adam J. Brunke, Canadian National Collection of Insects, Arachnids and Nematodes, Ottawa, Canada AVE Arthur V. Evans, Richmond, VA (also personal collection) CWH Curt W. Harden, VMNH, Martinsville, VA (also personal collection) DSC Donald S. Chandler, University of New Hampshire, Durham, New Hampshire (also personal collection) ERH E. Richard Hoebeke, Georgia Museum of Natural History, Athens, Georgia MKT Margaret K. Thayer, FMNH, Chicago, Illinois MLF Michael L. Ferro, Clemson University, Clemson, South Carolina Collections: AMNH American Museum of Natural History, New York, NY BMNH The Natural History Museum, London, Great Britain CNCI Canadian National Collection of Insects, Ottawa, Ontario, Canada CUAC Clemson University, Clemson, South Carolina DENH Department of Biological Sciences, University of New Hampshire, Durham, New Hampshire 42 BANISTERIA DFOC USFS Durham Field Office Forest Insect Collection, Durham, New Hampshire EGRC E. G. Riley collection, College Station, Texas EIUC Eastern Illinois University, Charleston, Illinois (as of 12/11/2019 part of FMNH) EJFC E. J. Ford collection (present location unknown) FMNH Field Museum of Natural History, Chicago, Illinois GWMP George Washington Memorial Parkway collection (property of U.S. National Park Service), McLean, VA MCZC Museum of Comparative Zoology, Harvard University, Cambridge, Massachusetts UMSP University of Minnesota, St. Paul, Minnesota USNM National Museum of Natural History, Washington, DC VMNH Virginia Museum of Natural History, Martinsville, VA VPIC Virginia Polytechnic Institute and State University, Blacksburg, VA Species are listed in alphabetical sequence by higher taxon, genus and species. Subgenera are indicated only when widely used in North America or when sometimes treated as genera, but do not affect the species sequence. Species that are believed to be adventive (“introduced”) in North America (from the Palaearctic region, with the exception of the Neotropical Anotylus insignitus) are indicated by a dagger (+). Those that are considered to be Holarctic (or even more broadly distributed) are indicated by an asterisk (*). New state or district records are boldfaced and marked with an exclamation point, and the source(s) of such records indicated in the “Source” column using the above abbreviations. LITERATURE CITED Blackwelder, R. E. 1973a. Checklist of the Staphylinidae of Canada, United States, Mexico, Central America and the West Indies. Family no. 15 (yellow version). North American Beetle Fauna Project; Biological Research Institute of America, Inc., Siena College, Loudonville, NY. 166 [as 165] pp. Blackwelder, R. E. 1973b. Checklist of the Scydmaenidae of Canada, United States, Mexico, Central America and the West Indies. Family no. 23 (red version). North American Beetle Fauna Project; Biological Research Institute of America, Inc., Latham, NY. 7 pp. NO. 53, 2019 Bousquet, Y., P. Bouchard, A. E. Davies, & D. S. Sikes. 2013. Checklist of Beetles (Coleoptera) of Canada and Alaska. Second edition. Pensoft Series Faunistica No. 109, Pensoft, Sofia-Moscow. 402 pp. Brown, J. W. 2008. Appendix. List of the invertebrates of Plummers Island, Maryland. /n J. W. Brown (ed.), The invertebrate fauna of Plummers Island, Maryland. Bulletin of the Biological Society of Washington 15: 192-226. Brunke, A., A. Newton, J. Klimaszewski, C. Majka, & S. Marshall. 2011. Staphylinidae of eastern Canada and adjacent United States. Key to subfamilies; Staphylininae: tribes and subtribes, and species of Staphylinina. Canadian Journal of Arthropod Identification 12: 1-110. Online’ version at <http://dx.doi.org/10.3752/cjai.2011.12> Last accessed 8 February 2019. Chandler, D. S. 1997. A catalog of the Coleoptera of America north of Mexico. Family: Pselaphidae. United States Department of Agriculture, Agriculture Handbook 529-31, Washington DC. 118 pp. Downie, N. M., & R. H. Arnett, Jr. 1996. The Beetles of Northeastern North America. Vols. 1, 2. Sandhill Crane Press, Gainesville, FL. 1,721 pp. Gusarov, V. I. 2003. A catalogue of the athetine species of America north of Mexico (Coleoptera, Staphylinidae, Aleocharinae, Athetini). <https://web.archive.org/web/ 20100613213828/http://www.nhm.ku.edu/ksem/peet/ catalogs/cataweb.htm>. Last updated 15 December 2003; last accessed 8 February 2019. Herman, L. H. 2001. Catalog of the Staphylinidae (Insecta: Coleoptera). 1758 to the end of the second millennium. Parts I-VII. Bulletin of the American Museum of Natural History 265: 14218 (in 7 vols.). Horn, G. H. 1868. Catalogue of Coleoptera from southwestern Virginia. Transactions of the American Entomological Society 2: 123-128. Klimaszewski, J., A. Brunke, V. Assing, D. W. Langor, A. F. Newton, C. Bourdon, G. Pelletier, R. P. Webster, L. Herman, L. Perdereau, A. Davies, A. Smetana, D. S. Chandler, C. Majka, & G. G. E. Scudder. 2013. Synopsis of adventive species of Coleoptera (Insecta) recorded from Canada. Part 2: Staphylinidae. Pensoft Series Faunistica No. 104, Pensoft, Sofia-Moscow. 360 pp. BRATTAIN ET AL.: ROVE BEETLES 43 Klimaszewski, J., R. P. Webster, D. W. Langor, A. Brunke, A. Davies, C. Bourdon, M. Labrecque, A. F. Newton, J.-A. Dorval, & J. H. Frank. 2018. Aleocharine rove beetles of eastern Canada (Coleoptera, Staphylinidae, | Aleocharinae): a glimpse of megadiversity. Springer-Verlag, Cham, Switzerland. 902 pp. Leng, C. W. 1920. Catalogue of the Coleoptera of America, North of Mexico. J. D. Sherman, Jr., Mount Vernon, NY. 470 pp. Lobl, I. 2018. Coleoptera: Staphylinidae: Scaphidiinae. In World Catalogue of Insects. Vol. 16. E. J. Brill, Leiden, The Netherlands. 418 pp. Moore, I., & E. F. Legner. 1975. A catalogue of the Staphylinidae of America North of Mexico (Coleoptera). University of California Division of Agricultural Sciences Special Publication 3015: 1-514. Newton, A. F. 2019. Staphyliniformia world catalog database (version Nov. 2018): Staphylinoidea, Hydrophiloidea, Synteliidae. Jn Roskov, Y., G. Ower et al. (eds.). Species 2000 & ITIS Catalogue of Life. Species 2000: Naturalis, Leiden, the Netherlands. www.catalogueoflife.org/col. Last accessed 8 February 2019. Newton, A. F., M. K. Thayer, J. S. Ashe, & D. S. Chandler. 2000. Staphylinidae Latreille, 1802. Pp. 272— 418 In R. H. Arnett, Jr. & M. C. Thomas (eds.), American Beetles, Vol. 1. Archostemata, Myxophaga, Adephaga, Polyphaga: Staphyliniformia. CRC Press, Boca Raton, FL. 443 pp. O’Keefe, S. T. 2000. Scydmaenidae Leach, 1815. Pp. 259-267 In R. H. Arnett, Jr. & M. C. Thomas (eds.), American Beetles, Vol. 1. Archostemata, Myxophaga, Adephaga, Polyphaga: Staphyliniformia. CRC Press, Boca Raton, FL. 443 pp. Ulke, H. 1902. A list of the beetles of the District of Columbia. Proceedings of the United States National Museum 25 (1275): 1-57. NO. 53, 2019 BANISTERIA 44 HM) u! HM9-VWA dWMD U! Hdus-VA dWMD U! Hdus-VA dWMD U! Hdus-VA dWMD U! Hdus-VA HM) u! HM9-VWA HNWA U! HM9-VA 301NO0S GW aw aw od 3d od (pr8T ‘ewWleysjaW *J"4) e7sepow (ejo.4awiq) ejayjy LOEST ‘WieyjeuUeW|) Biazdouejseo (ejayjy) e]eyIy OT6T ‘Aasep exouue (ejopnasd) ejJayiVy (6E8T ‘UOsyoH3) enwee (ejay;y) EIOYIY I(ZO8T ‘JSHOUUSAeID) S/eUe CYISILUY (6€8T ‘UOSUdZ) ey/puora/ CUOJOIOY (OT6T ‘Aase>) e7e/ND eUOIOIOY (6E8T ‘UOSUDA) E/NISNUAA B1BUOLOW/ (OT6T ‘AaseD) eYAqis eUly2@aIGO/eL (606T ‘Joneyuteg) sde20edo eulyjIe/qGO/eCH €00Z ‘AoJesNy Maseo esnjuoy 906T ‘AaseD syuooijas eluojedy *fE8T ‘Aes CLA (B/EYI0ICOD) e1eYyI0a/Y (€6gT ‘Aase)) sLAUAARAaINIS (eseYIOUBX) e1eYIOAY OT6T ‘Aagjyojelg sedugns (eseyooje7) eseyI0a/p b86T ‘DISMAZSELUI[Y SLE/NIO (e/eYDOYIF) e1eyI0a/ ZEST ‘Aes eILsry (eleyroa/y) e1eyr0a/V (c6gt ‘Aasey) ebnyon (esey20yrz) e1eyr0a/V (€S8T ‘UIPIEW) S/e/O] (EJoUe/duz) e1eysoa/Y LZO8T ‘JsHOYUSARID epey (eveYI0a//) eley0a/y TO6T ‘saneywag s/o moe (eseyI0a/p/) eseyI0a/P 1ZO8T ‘JSHOYUSAeID e7ewury (eseYIOUAaX) e/eYIOI/Y M(LLLT '8Z805) eynyina (eleyso0a/y/) eleyr0ajy CPST ‘WidayjauUep| sUUediauejse) (eJCYIOUBX) e1eYD0A/Y L107 ‘ayxUNIg g JaISGaM\ ‘DJSMAZSEUUI) BEJOLLELUIBUEISED (CJEYIOUAX) CJCYIOI/Y ZOST “JSIOYUdARID Eyenzewuig (eseyI0sdo7) eseyI0a/V awey eUnauyy eUnauyy eUNauyy eUnauyy eUnauyy eUnauyy eUNauyy PULIEYDOR|V PULIEYDOR|V PULIEYDOS|V PULIEYDOR|V PULIEYDOR|V PULIEYDOR|V PULIEYDOR|V PULIEYDOR|V PULIEYDOR|V PULIEUDOR|V PULIEYDOR|V PULIEYDOR|V PULIEYDOR|V PULIEYDOS|V aqinqns JUnaLpY jUnaLaY JUnaLDY JUnaLDY IUnaLDY jUnaLDY JUnaLDY jUnaLpY JUnaLDY JUnaLpY JUnaLDY Juneau IULEYDOR|Y IULEYDOR|V IULEYDOR|Y JULEYDOa|V IULEYDOS|Y IULEYDOR|Y IULEYDOR|Y IULEYDOS|V IULYDOR|VY IULEYDOR|Y IULEYDOR|Y IULEYDOR|V IULEYDOR|Y IULLEYDOR|V oquL aquyedns seULIEYOR|V seULIeY ORV seULIeYyoa|y seULIeYOR|V seULIeY ORV seULIeY ORV seULIEYDOI|V seULIEYDOI|V seULIEYDOI|V seULIEYDOI|V seULIEYDOI|V seULIEYDOI|V seULIEYDOI|V seULIEYDOI|V seULIEYDOI|V seULIeYOR|Vy seUuLeYyoa|y seULIeYOR|V seULIeY ORV seULIEYDOIIV seULIEYDOI|V seULIEYDOI|V seULIEYDOI|V seULIEYDOI|V seULIEYDOI|V seULIEYDOIIV Ajiueyqns XIGNUddV 45 BRATTAIN ET AL.: ROVE BEETLES VA VA od od +(9G8T ‘UOSWOY| *5°D) e72aSU/ BJOUOIO/Y (906T ‘Aeses) sy/9e/B snjeYdaIOIALLLAp/ x(6E8T “UOSUDHF) SULIDUOD SN/EYABIOIOULIAP/ (998T ‘2}U0D27 “1'C) sMe/NBUID SN/EYABIOIBLLIA| (OZ6T ‘UeWION) sdea7e/ eLiBess/7 6E8T ‘UOSUDHA epeassip eubejey LZERT ‘UOsYoLA esae eubeyey I(ZO8T ‘JStOYUSAPIO) e/NDSgo el/ep10D (998T ‘2]U099) “1'[) sdeaupenb e1odosoquog (998T ‘a]U0D27] “1'C) sveyaznas snuopoa/y (998T ‘2}U0D27 “1'C) snyed sniopos/y eeet ‘Aes snjegojiq sniopoa/y Q06T ‘AaseD EX/UIBIIA eSNIYOU] 906T ‘AaSe)-/O/ODLIEA ESNIYOL] C68T ‘Aasey enysngos esniyotl] 906T ‘AaseD e7//od esniyoU] 9Q06T ‘AeseD E7NSUIL ESNIYOLL C68T ‘Aase) eIEALUOD BSNIYIL] (ZO8T ‘JSJOYUSAID) eO/YDID eSNYJAIS L(OERT ‘WIayoUUeL|) S/UUadIpIAy eIdO.WILUaYeaN (6€8T ‘Uosyst3) e/oazny eyA20~y (OT6T ‘Aase>) xapne? eJAYJEIOIP/ (998T ‘a}U0De7 “1'C) esnigo sueinbue essojbodAT 906T ‘Aase epiony ejowWoINF ZO6T ‘Janeyusag esorLyUAA (elgoe/y) ejay]y Q86T ‘Ped B PsmazseUl|y E/Ydo/bo04 (ej0.HELUIG) EIEYIY OT6T ‘Aesep euexa) BJO) OT6T ‘AeseD e/naHal eJOYIY TT6T ‘AaseD esa EJAY] (OT6T ‘Aases) epeuosiad (e}0/ANON) BJAY]Y ZO6T ‘Janeyuwag exuerjAsuuad (eJ0palolp|) e}aYIP eUnauyy eUnauyy eUnauyy eUnauyy eUnauyY eUnauyy eUnauyy eUnauyy eunalyy eUNauyY eUnauyy eUnauyy eUnauyy eUnauyy eUnauyy eUnauyy eUnauyy eUnauyy eUnauyy [UIGSOe5 1ubeje4 jubeje4 jubeje4 juibeje4 jubeje4 juilibeje4 juiibeje4 jubeje4 jubeje4 jubeje4 juiibeje4 lUOUW lUOUW |UOUIW |UOUW lUOUIW lUnOUW |UOUW lUOUW lUOUYW lUOUYW lUOUIW lUOUW |UOUIW lUOUW lUOUW lUOUW lUOUW lUOUYW |UOUW seULIeY ORV seULIeY ORV seUuLIeYoR|y seULIeY ORV seULIeY ORV seULIeYOR|y seULIeY ORV seULIeYOR|y seULIeY ORV seuLIeYyoa|y seULIeYOR|V seULIeY ORV seULIeYy ORV seULIeY ORV seULIeYOR|V seULIeY ORV seULIeYOR|y seULIeYOR Vy seULIeY ORV seUuLeYyoa|y seUuLeYyoa|y seULIeYOR Vy seULIeYOR|y seULIeYOR Vy seULIeYoR|y seULIeY ORV seULIeY ORV seUuLeYy oR |y seULIeYOa|y seULIeY ORV seULIEYDOI|V NO. 53, 2019 BANISTERIA 46 VA VA VA VA dW aw oe Od od (GE8T ‘WOSYDIF) SI/LUISSIPD PJOJBUEYd Q06T ‘Aasea eULZIA eUaeYdoIAD TG6T ‘SHAABVS SUB/NLUIS PUBCYAOIAD TS6T ‘SHAABdS SAaD/OUeU CUBCYAOIAD 906T ‘Aase e7ego/ BEUaeYAdOIAD 906T ‘AaeseD e/njaey euaeYydoIAD TS6I ‘SdaAvas sBIeggny euaeydo‘AD Q06T ‘Aase) sH/ODIA/IB euaeydosAD 9Q06T ‘AaseD sy//ODDSnyY BEUAYAOIAD bPbST ‘JawWlaysjay| "34 S/LWODNeY PUALYAOIAD Q06T ‘AdSeD SLUAA/IIUOD BUBeYAOIAD TS6T ‘SdaAv—as Vapjamyoeq eueeydosAD LOEST ‘Wiaysouuep sue EUAeYAOIAD (6E8T “UOSUDZ) e905 EJOJNUNF (6E8T ‘UOSUIHZ) E/NISNLOD EJOJIOMUNF (6E8T ‘UOSUDINF) EJE/NICLUILY IGOJOINAd C6gT ‘Aaser e2edo (eesidoxopy) esnjda7 OT6T ‘Aagynielg suebayja (eyesidoxopy) esnjda7 (906T ‘Aase)) enjeiquo (esnyyj) esnjdeT C68T ‘Aasey syjooinaig (esnjdAionz) esnjdeT GT6I ‘AaseD sdezjpuelb esnulwA5 6/6T ‘DISMAZSCUUIPY ELUE/LIBIIA SISAOLIAG 6Z6T ‘DISMAZSeUUIPY ELUIPEY SIsdoulag TT6T ‘Aases epides sisdoulag ZS8T ‘Aeesy snuequawe sisdoulag (ZS8T ‘ZWee1y) sapiouse|/AWW sisdouipy Z86T “ISMAazZSeUUI}y E7ep/dsnd sisdouipy 7002 ‘AOJesny euabiyoejedde equsoay (906T ‘Janeyusag) e7e7uap ejosez x(ZEST ‘Suaydays) suayiojns ejounojojyY euluaeydosA5y euluaeydosAy euluseydosAy euluaeydosAy euluaeydosAy euluaeydosAy euluaeydosAy euluaeydosA5y euluaeydosA5 euluaeydosAy euluaeydosAy euluaeydosA5 euluaeydosA5y euluaeydosAy euluaeydosA5 PULEYDO}|Og PULEYD0}|Og PULIEYDO}|Og PULEYDO}|Og PULIEYDO}|Og |UO|eWOH JURO|EWOH JURO|EWOH JURO|EWOH JURO|EWOH JURO|EWOH JURO|EWOH JURO|EWOH JURO|EWOH JURO|EWOH JURO|EWOH JURO|EWOH JURO|EWOH JURO|EWOH JURO|EWOH JURO|EWOH JURO|EWOH JURO|EWOH JURO|EWOH JURO|EWOH IUISNUWAS IUISNUWAS IUISNUWAS IUISNUWAS IUISNUWAS IUISNUWAS IUISNUWAS 1UIGIISO85) |UIGIISO85) JUIqISOa5) seULIEYDOI|V seULIeYyoa|y seULIeYOR|V seULIeY ORV seULIeYOR|V seULIeY ORV seULIeYOa|y seULIeY ORV seULIeYOR|V seULIeYoa|y seULIeY ORV seULIeY ORV seULIeY ORV seULIeYOR|y seULIeYOR|V seULIeYOR|y seULIeY ORV seULIeY oa |y seULIeYoa|y seUuLeYyoa|y seULIeYOR|V seULIeY ORV seULIeYOR|V seULIeYOR|V seULIeYOR|V seULIeY ORV seULIeYy oa |y seULIeYOa|y seULIeY ORV seULIEYDOI|V 47 BRATTAIN ET AL.: ROVE BEETLES HM) u! HM9-VWA iVA VA VA VA VA VA GW GW GW Od oe 3d Od 3d €68T ‘Aasen esadse esnhoQ (6st ‘Aesea) sLauannsnbue elAboixaq (TZ6T ‘seAue4) Vaqueqg essojbojdey L(OEST ‘WiaysouUeL|) syVe/nyns eaeseje1 (p68T ‘UURLUSeM) SI//Gap XeIOYINT (9991 ‘a\U0 27 *7"C) esuedxa esnzaq TT6T ‘Aesep enpiooid euael/Ap/ Z86T ‘PISMEZSELUIP /LUOJEMejOd euae//Ap/ Z86T ‘DISMAZSEWIP EryYSeysey eUae//AW/ *€S8T ‘eer eyersryul EUae|/AW OZ6T ‘UEWION B7eaUND euae//AW (€98T ‘a}U0De7 “1'[) sNaIe}sa} sndAIeUy (9981 ‘a3U0D27 “1'C) SIO (SeIAZ) SeIAZ (e6gT ‘Aasep) sayueld (seiAZ) seIAZ (€68T ‘Aased) e7e2U0/ e//Ad (‘AaseD) esourbyed el/ad 6S6I ‘SIOABIS SILLIODINAIG EIPIUAXOPIF 710z ‘ebpeipla sayse eyisnig (998T ‘Jay/EM) exoya/ eEsnpOouaX (€98T ‘aJU0De7 “1'C) BAe2 esnpousXx 18S8T ‘seQemuasaly oy/uNnd e060 OT6T ‘AaseD SeUI/O/E CLIDUR}E/d (6E8T ‘UOSYDF) BADeYIS E/OJII~ (pPpST ‘Jawlaysjeay) *3°4) s/e/a7e/ (eELipue;dopH) eLypuedoy OT6T ‘Aasey sLaUAaAINae/ (eELipueUaYLy) eLipue/doyy (OZ6T “UPWION) syjoriAae) (WaJDNWOYAOT) eLjpueldoH 686T ‘181UaD /P/SMAZSELUI/Y (JAJINWIOYAOT) eLpue/doy{ C68T ‘Aasey o7/de? eJ0INIIAY | L(OTST ‘eyual|Ad) eued ejo/ewopy (ZE8T ‘AeS) EFEISE EJOJBUCYd PUIDOdAXO PUIPOdAXO eUIMO|HOsI[)) eUIMO|HosI[)) eulpjeuig eulpjeuig PUIIUOPALLA|,| PUIIUOPALWA|\| PUIIUOPALWA|\| PUIIUOPALWAI\| PUIIUOPSLLA|\| PUIIUOPALWA|\| PUISNYDSWO7] PUISNYD@WOT PUILPUe}e}d PUILPUe}e}dq eullipue|doy eullupue|doy eulLipue|doH eulLipue|doy PUIO|ELUOH PUIJO|ELWOH euluaeydosA5 IUIPOdAXO lUIPOdAXO lUIPOdAXOQ lUIPOdAXO |UIPOdAXO IUIPOdAXO jujUse||AW JUJUae| Aj] JUJUSe| Aly] JUJUSe| Aly] JUJUSe| Aly] lULOdosd|\| IUISNYDOWO] IUISNYDSWO] IUISNYDSWO] IUISNYDSWO] |UISNYDAWO7 JUISNYDSWO7 JUISNYDSWO7 IUISNYDSWO7 lunYydADOdAH lulLpue|doy julLpue|doH lullipue|doy lulLpue|doH julLpue|doy lulLpue|doy [UIJO]/EWOH |UIJO|EWOH JUO|EWOH seULIeY ORV seULIeYyoOa|y seUuLIeYyoa|y seULIeYOI Vy seULIEYDOI|V seULIeYOR Vy seULIeYOR|V seULeYyoI|y seULIeYOR|V seULIeY ORV seULIeYOa|y seULIeYOR|y seULIeYOa|y seULeYoa|y seUuLeYoa|y seULIeY ORV seULIEYDOI|V seuLeYyoa|y seULIeYOR|V seUuLeYoa|y seULIeYOR|V seULIeY ORV seULIeYOR|V seULIeY ORV seULIeYOR|V seUuLIeYy ORV seUuLIeYoa|y seULIeYOR|y seULIeY Oa Vy seULIEYDOI|V NO. 53, 2019 BANISTERIA 48 HM) u! HM9-VA HM) u! HM9-VWA AAV Ul SAV-VA VA VA VA VA aw GW P88T ‘AdSeD SI//LLIS SNYBYISBENT b8gT ‘Aasen smjejuNd snjayjsaenz P8ET ‘Aase sninpuNW snjayjsaenz p8gT ‘Aasey sIULIadLI snjayjsaenz OP8T ‘UOSUDHA snuequeWwe snjayjysaenz LS8T ‘AysiNnYyDs}OW SNPAIU snydepz bZ6T ‘ZuUINd snUeQVEWwe snydepF Q66T ‘AWEIeD 1g |GQ]4O/0DU0D snuadAseq Z88T ‘UIOH sisuauoler sniedAseq 4T98T ‘UOSWOY] *5°D sadiney ejIa/GO/EH 906T ‘AeseD e7ajosgo esnAyoe | CQQT ‘a}U0De7 “71'C eLuyjoe/B esnAyre 1 Ggst ‘Aasep e7azey esnAyoe | EQ8T ‘a}U0D2] “TC SY/OIINeD esNAyre | 906T ‘Aasep eueuVawe esnkyre 1 (c9gT ‘ayU0De7 “1°C) eye/biu eyadAuH (€9gT ‘equoDe7 71°C) Baywyeg ejadhuy C68T ‘AdSe> SCLUOIE] ESNIE|d 6E8T ‘UOSYdZ eedsap esnre/d Z00Z ‘EXLEW 8 PISMAZSELUI|Y JELLO EANF LG8T ‘Z}ee41 SNSO/IA SALLLIAJO/IYd ZS8T ‘Zee1y SNIIUEA/ASUAd SALLLIAJO/IYd LG8T ‘ZIECID| MSYINY SALIAIO//Yd LET ‘SIBAVAS SI//COJIGNI SAULIAJO/IYd 906T ‘AeseD siaeqns e1odoa0/Yd x(ZO8T ‘JSIOYUSAPI) SYEIZIOI e1OdO0a0/Yd 6E8T ‘UOsUouy e7ejnbes epodAxOQ 906T ‘Aesea enbyqgo epodAxoC €68T ‘Assen sdeaubiu epodAxCQ 906T ‘AaseD e27aLUIL EpOdAxO PULOdOaO|Ud PULOdOaO|Ud PUIPOdAXO PUIPOdAXO PUIPOdAXO PUIDOdAXO luljey}SeeNy lujeyyseeNy luljey}SeeNy luljeyyseeny luljeyyseeny luljeyysoeny lujeyyseeny JUL9DIXe | luisnAyoe | luisnAyoe | luisnAyoe | luisnAyoe | luisnAyoe | luisnAyoe | luisnAyoe | IUISNIL|d IUISNIE|d IUISNIL|d JUNILLI}O|IUd JUNILLIJO|IUd JUNILLIA}O|1Ud JUIJLLLI3}O|1Ud |UIPOdAXO lUIPOdAXO IUIPOdAXQ lUIPOdAXO IUIPOdAXOQ lUIPOdAXO seuljoy}soenz aseuljeYy}seeny seuljoyjseenz seuljoYy}seeny seuljey}seeny aseuljoy}seeny seuljoyj}seeny aeuliadAseg aeuliadAseg seuLeYyoo[y seuLIeYyoo|V seuLeYooly seuLeYyoo|y seuLeYypooly seuLeYyooiy seuLeYypooly seULIEYDOIIV seULIEyDOI|V seULIEYDOI|V seULIEYDOI|V seULIEYDOI|V seULIEYDOR|V seULIEYDOI|V seuLeYyoa|y seULIeYOR|V seULIeYy ORV seUuLeYyoa|y seULIeYOR Vy seULIeY ORV seULIEYDOI|V 49 BRATTAIN ET AL.: ROVE BEETLES HNWS U! DIW-VA WNSN u! DIW-VA HNWA ul! DIW-VA S4SN U! DIW-9d HNWS U! DIW-VA iVA VA VA VA VA VA VA VA VA GW GW QW GW GW GW Od 3d od Od od od C68T ‘AeseD auuadupenb winjeuC 8/8T ‘Jaane Lwinquo wNniewuo (S/8T ‘Jeane4) e7ewey eaesejedey (S68T ‘UO ILWEH) sNpIeYy snjeyded0uizie) (QZ8T ‘jeAne4) esnyip eysojai0y (8/8T ‘jeAne4) soy Linuajeydsnz (SZ8T “JeAne4) Lnxeduo) wnsaeydsnz PlOZ ‘Meuez sisueuyosen winsejeydsnz PRST ‘ejU0De7 “T'( SO DIsselD snyIapNF (€98T ‘ajUODeT “T'[) sNjJejOU snujayAaz (€98T ‘a}U0De7 “1'C) 5/7e73Nb snuyjayaz CQQT ‘9JU0D2] “1'( SMELIS SNUIAPOUOBL | OP8T ‘UOSYdUF LwnzIajgGo LniydojO «(OTST ‘Jeyual|AD) aytuisuos wunsydojO E98T ‘A}U0De7 “TC seayjed erajsaT (c6gt ‘Aase)) E/NJEIGUD ENAISIT 688T ‘leAney SMpILYS SNIILUOIpOAD (€ZET ‘AeS) sneuUTUG sNoILUOJpOay ZS8T ‘8}U0D27 “TC SNpAIU snuIyelg 8/8T ‘jaaney /Zemyos Lunipadiy (Ob8T ‘UOsysII3) LuNpIplos WiniqoyWjUy OP8T ‘UOSsUdHy eyeuLiesqns ejopioy x(Z6ZT ‘Sniouqe4) e7eua.o ejopioy CORT ‘8}U0DET “TC SNPEIQUO SN/dado.Ipy S96T ‘||eqdwey /UMaIg SsN/dedo/ip (2581 ‘Aysinuas}op|) sedyns snuidojebay (ZO8T ‘JS1oyUdAeI) sme/aeD snuldojebay/ 998T ‘38]U0D97 1'¢ Ssdea2UNd sniuesIOJINS eullydAo7 eullydAio7 eullyudAio7 JUII|ELUO JUII|ELUO JUII|ELUQ 1UII|ELUQ 1UII|ELUQ luLajeydsny luLajeydsny luLiajeydsny juNydAoD juNydAoD juNydAoD juIbeydouuy juibeydouuy juibeydoujuy luibeydoujuy luibeydoujuy luibeydouuy juibeydouuy juibeydouuy juibeydouuy juibeydouuy luibeydouuy juibeydouuy JUNULIIOJNS SEUIIJEWCO SEUIIJEWO SEUIIJELWUCO SEUIITEWO DEUII|EWIO DEUIITEWIO SEUIITEWIO SEUII|EWO DEUIITEWO DEUII|ELIO SEUII|EWIO SEUIITEWIO SEUII|EWIO DEUIITEWIO SEUIITEWIO SEUIIJEWO SEUIITEWIO DEUII|ELUO SEUIITEWIO DEUII|ELUIO SEUIITEWIO DEUII|ELUO SEUIIJEWO aeuljdados3iW aeuljdadossiw aeulipisdojebaw aeulipisdojebow seuljeYyysoeNny NO. 53, 2019 BANISTERIA 50 dWM9-VA dWM9-VA dWM9-VA WNSN Ul DIW “dWM9 -VA ‘dSWN Ul DIW-dW WNSN u! DIN-VA HNWA ‘dWM9 UI DIN-VA WNSN Ul DIW ‘dWM9D “VA iVA iVA iVA iVA dW aw idW od od od Od od od (€9gT ‘a]U0De7 “1'C) smenzewanbuinb snuodAxopneasd (POST ‘Jeane4) syeydin[0 snsodAxopnasd (ZO8T “JsJOYyUdAeID) syesaze/ snuodAxopnas” ZOST “JSIOYUSARID SMeWIA snuOdAxO TEst ‘Aes snaibAjs snuodkxoQ €QQT ‘8]U0De7 “TC SlUUadYyrU snIodKxE OO8T ‘JS1OYUdAeI vole snuodAxOQ LL8T ‘83U0De7 “TC sNpidey snsodAxE ZOST “JSIOYUDACID S/C/IOLUAY SI/CJOLUAY SNIOAAXE (OP8T ‘UOSYDF) S/e7SOD SNIOYAODEJOY/ (9981 ‘a}U0D27 “T'C) EpNeWNae/ eIGUOld (€98T ‘a]U0D27 “1'[) sUaDsayny SNUIASI/IAEID (OP8T ‘UOsUdIIZ) sNNBixe snurdsyinep (908T ‘JSuoYUaAeID) sadyey ELOSO/OW GO6T ‘JeANe4 SIAIAAYIEIG SNYIOAO/OLY (€ggt ‘ajU0De7 “1'C) eyavbiu eypseuay (SggT ‘a]U0De7 “7°1) epI/ed sisna/z €007 ‘saAeul Wayns SNLUOIpO/AX I(ZO8T ‘Wweysiep|) SMUUIIUOD SNIOIpO/AX (SZ8T ‘JaAne4) supUeAGoUNd edaspol/AYd (SZ8T ‘jeAne4) esosaluny edeaspol/Ayd (2ST ‘UIPEW) SI/02/SAL/ SNUIOUOBOIYd 8/8T ‘jaane4 winjzey (0je| NSuas) Lwny/eLUC 4(68ZT ‘|IMAed) venAL LuN/eLUC OP8T ‘UOsUoH Lunpuede/ WNyeUuCQ euloydooesou | eUIUICSI|IAe|D BUIUICSI|IAe|D eUIUICSIIAe|D luUUOYdode40U | luUUOYdode40U | luULOYdode40U | luUOYdode10U | 1UILOSC 1UILIOSC luluisna|y luluisna|y 1UII|ELUQ 1UII|ELUQ JUII|ELUO JUI||ELUO JUII|ELUO JUI||ELUO JUII|ELUQ JUI||ELUO aeUulLIOdAXO BeULIOdAXO seUuLIOdAXO aeULIOdAXO deULIOdAXO BeULIOdAXO aeUuLIOdAXO aeUuLIOdAXO aeUulLIOdAxO SeUILIOSO SEUILIOSO SEUILIOSO SeUILIOSO SEUILIOSO SeUILIOSO DeUILIOSO SeUILIOSO SeULI|EWIC SEULIJEWUIC SEUIIJEWO SEUIIJEWO SEUIIJEWO SEUIIJEWCO SEUIIJEWCO SEUIIJEWO 1 a) BRATTAIN ET AL.: ROVE BEETLES dWM9-VA dWM9-VA dWM9-VA iVA VA VA GW od Od od Od LOO8T ‘ISIOYUdARIN SN7AQ/NIS sNjayAxO OP8T ‘UOsYyoHy snzjued/Asuad snjajzAxOQ 1(ZO8T ‘weysiep|) snzeanbe snjazAxo C68T ‘AdSeD SNSNUAALI SN/AIAXO OP8T “‘UOSUD SILUN/OIUI SNjaJAXO (TEst ‘AeS) syjooueeyds snjjaz0dy (c6gt ‘Aasey) snzzedsns sn/Ajouy KGZZT ‘sniduqe4) snsobn snjAjouy (ZZ8T ‘eqU0De7 “1'[) snUisNIe|/d snjAjouy (OP8T ‘UOSydy) snueUu sn/Ajouy L(908T ‘JSIOYUSACID) SNZUBISUI Ssn/AjOUY (OP8T ‘Uuosydy) snnbixa sn/Ajouy (c6gt ‘Aase)) snsuap sn/Ajouy LL8T ‘aquodeT “1'(. 727 snipalg LL8T ‘aU0De7 “1'( SMENUIS SNIpAlg COST ‘a}U0D27 “T' sNaW/bnuayiwes snipag OPST ‘UOSUDHZ snsoulbiqnu snipalg LL8T ‘2\U0DeT “TC snuussyejouNd snipag OP8T ‘UOsYsA snyyod snipea/g 6T6T ‘Ile4 snaiyqaperyd snipalg E98T ‘a}U0De7] “TC sN/NIedO snipalg E98T ‘AUODST “TL SI/ODIPHIU Snipa 688T ‘Aese 5172a/bau snipa/g OP8T ‘UOSUDHA SsUe;NngpueW snipag (Test ‘Aes) snjeuibyewea snipa/g (TEgt ‘Aes) smpepsoo snipalg LL8T ‘AU0D27 “TL SNSNyUOd Snipa E98T ‘aUODeT “TL syeseg snipalg 688T ‘AaseD sy//uisse snIpa/g E98T ‘aUODe] “TL Sve;NuUe Snipa €Q8T ‘a]uDe7 “Tr syeue snipalg |Ul[2}AXO |Ul|2}AXO 1Ul]2}AXO |Ul|2}AXO |Ul[2}AXO |Ul|2}AXO |Ul]2}AXO |Ul|2}AXO |Ul[2}AXO 1Ul[2}AXO |Ul]2}AXO |Ul[2}AXO |Ul[2}AXO julIlpald lulIpald julIlpald julIpald jullpald lulIpald lulIlpald lulIlpald lulIlpald lulIlpald lulIlpald lulIlpald julIpald julIlpald lulIlpald lulIlpald lulIlpald julIpa|d aeuljayAxO aeuljayAxO aeuljayAxO aeuljayAxO aeuljayAxO aeuljayAxO aeuljayAxO aeuljayAxo aeuljayAxO aeuljayAxO aeuljayAxO aeUuljazyAXO aeuljayAxO aeUljayAXO aeuljayAxO aeUuljayAXO aeuljayAxO aeuljayAxO aeuljayAxO aeUuljayAXO aeuljayAxO aeuljayAxoO aeuljayAxoO aeuljoyAxO aeuljayAxO aeuljayAxO aeuljayAxO aeuljazAXO aeuljayAxO deUljaJAXO aeuljayAxO NO. 53, 2019 BANISTERIA N a) dWM9-VA dWM9-VA dWM9-VA dWM9-VA iVA iVA VA GW GW Od (O88T ‘2}U0D87 “T'[) enbiqwe enpjejopnasd (ZO8T ‘JssouUaAeI) wnjNnosnibuog] wniyjesgoT (T86T ‘sno.nem) wnjeuibvewa wniyjesgo7 (OP8T ‘UOSYDLZ) a/e/02 WuNijJe/gOT (SO6T ‘AeSeD) LUNDIUIBIA LUNIGOIYJe7 (So6T ‘Aase>) wnzedsns wnigosyje7 EBT ‘9]UODO] “Tf /LLs LUNIGOIYJET (SO6T ‘AeSeD) LNJEWIOU! LUNIGOIYJeT OSst ‘Aes winjgeuue WNIgosyjeT (998T ‘AaSe)) SILUODINAIG SAISEIYIF (OP8T ‘UOSUda) snxiaid snuajsy (O€8T ‘WilayjauUeL]) s7/NISN/BUO/ SnuaISyY (SO6T ‘Aase>) sde2Dsry snuajsy (ZZ8T ‘unsny) siuuadinaig snuajsy (Ezet ‘Aes) smejoulq snuajsy (SO6T ‘Aese)) snuequeWe snuajsy (688T ‘AaSe)) SNUIAIOD SNLIOJPOUIY 1 (ZZ8T ‘8}U0D27 “7'C) 4afove SNWOIPOUIyL (LI6T ‘Haqnyups 1g Janeyusag) SNUeIAEWe SNLWIOSPOUIY{ LL8T ‘9\U0D2] “1'[ SNPELIQUIY STIGOUIL (ZZ8T ‘a4Uu0De7 *7'C) SiloyiuN snumjedsez 4(6E8T “UOSYoLF) SsyAgns snujadie (688T ‘AeseD) snjaids snujadieD L(O98T ‘AYSINYDS}OW!) SHE/NAL SNUNjadeD (Test ‘Aes) smjejoundupenb snumjadiez L(ZO8T ‘JsHOYUaAeIN) sNyisNnd snumjadse? L(OEST ‘WiayaUUeL|) syOe/E snumjadie? (OP8T ‘UOSsUdA) sadiajry snuujadiez (688 ‘Aesed) SYOYYIP SMLUIjadleD IPERT ‘suaydays snzeouiig snumjadie) ObST ‘UOsUdIIa snue quae snyjajysied PUIIGOIYIET eUIIGOIYIET PUIIGOIYE7 PUIIGOIYIE7 eUIIGOIUE7 PULIGOIYE7 PUIIGOIYIE7 eUIIGOIYE7 eUIIGOIYIE7 PULIA}SeIYDI PulUa]sY eUlUaSY eulua]sy/ eulua]sy/ eulud]sy/ eulua}sly 1UNIGO4Uye 7 1UNIGOIUe 7 1UNGOIUYe) 1UNIGOIUe) 1UNIqO41UIe 7 1UNIGOIUIe) 1UNIqOIUYe) 1UNGO4UJe) 1UNIGO4Uye) 1UNIGO4Ue 7 1UNIqOIUe 7 1UNIGO4UIe 7 1UNGO4UIe) 1UNIGOIUYe) 1UNIqO4Ue 7 1UNGOIUJe 7 1UIIQOUIY | 1UIIQOUIY | 1UIIQOUIY | IUIIQOUIY | 1UIIGOUIY | 1UNIQOUIYL [UNIQOUIY | 1UIIGOUIY | 1UIIQOUIY | 1UIIGOUIY | 1UIIQOUIY | IUNIQOUIY | (UIIQOUIY | 1UNIGOUIY | 1U!|2}AXQ aeuLlopsed aeulLlapeed aeuLlapsed aeulLlopeed aeullapeed deulLlapsed aeuLlapeed deuLlapsed aeullapsed aeullapeed aeuLlopsed aeulLlapeed aeullapeed seuLlapsed aeuLlapsed aeuLlapsed aeuljayAxO deUulja}AxO aeulja}AxO deUulj[a}AxO deUulja}AxO aeUulja}AxO aeulja}AxoO aeuljayAxO aeuljayAxO aeuljayAxO BeUI][a}AXO deUI[a}AxO deUulja}AxO aeulja}AxO aeuljayAxO 53 BRATTAIN ET AL.: ROVE BEETLES HNWA ul Hua-WA HMO +dWM9 ul Hua-WA HM) u! HM9-VWA HNWA ul HD ‘dWMD “VA dWM9-VA dWM9-VA VA VA Od od Od od Od Od od (OggT ‘2]U0De7 “1'C) smeueg snbny (Op8T ‘uosysiy) suenbue snjbny (prST ‘JOWIaYs|aW “J'4) SASLA SNIINSNF (eggq ‘aqu0De7 “1'0) s72ed0 snaedo2s (COST ‘2}U0De7 “1'L) SNPAIU snaedo2s OPST ‘UOsUudHy snnbixa snaedons Go6T ‘Aasep smeayap snaedors (O8gT ‘a]U0De7 “7"¢) ~abQuap sTuC (TEST ‘AeS) snJUAaNYUOI SNIUNS (OP8T ‘UOSYDF) SI/IXA SLIEYIOIIS GOG6T ‘AdSED SISUBLIOJED SLIEYIOIIS I(ZO8T ‘JSAOYUSACIN) CADEIYIO SLIEYIOY}T (Sso6T ‘AeseD) sedyny uopawodky (SO6T ‘Aase>) snysnbue uopawodsy (ZO8T ‘JSIOYUSAPI) SNUIDIZIOD SNYAIOLIOUAYIY GO6T ‘Aasea sn2IWa] snedoyeja | GO6T ‘AaseD sIUUadUGns snadoyej}a | (O88T ‘a}U0D27 “1'C) s7/NPYIU snadoye}a | (€9gT ‘28]U09e7 “1'() “abu snadoyeja] GO6T ‘Aase) snuepLuOY snadoyleja (€98T ‘a]U0D27 *1'f) sue/nbue snaedoyeja / (O88T ‘2}U0987 “T'[) s/equaA elyjejopnasd (Oggt ‘8}U0987 “T'L) enpyjed esyjejopnasd (So6T ‘AeseD) suajal esyjzejopnasd (SO6T ‘AeseD) sisveM/Y EljejopNasd (Ost ‘AeS) e7eipiup enyejopnasd (€88T ‘4dIAIANG) eueqvawe e1yjejopneasd PUIDIINS PUIDIINS PUIDI|NS eulaedoos eulaedoos eulaedoos eulaedoos eulaedoos PUIUOpEe| PUIUOpe| PUIUOpe| PUIUOpaL| PUIUOpe| PUIUOPAaL)| PUIUOpEe| PUIIGOIUIE7 PUIIGOIYIe7 PUIIGOIUIe7 PUIIGOIUIe7 BUIIGOIUIE7 PUIIGOIYIe7 PUIIGOIUIe7 PUIIGOIUIe7 PUIIGOIUIe7 BUIIGOIYIE7 PUIIGOIUIe7 PULIGOIYIET JUNGOsUeT JUNGOsUeT JUNGOsUeT JUNGOsUeT JUNGOsUeT JUNGOsUeT JUNGOsUeT JUNGOsUeT JUNGOsUeT JUNGOsUeT JUNGOsUeT JUNGOsUeT JUNGOsUeT JUNGOseT JUNGOseT JUNGOsUeT JUNGOsUeT JUNGOsUeT JUNGOsUeT JUNGOsUeT JUNGOsUeT JUNGOsUeT JUNGOsUeT JUNGOsUeT JUNGOseT JUNGOsUeT JUNGOsUET seUuLIapsed seUuLlapeed seuLlapeed seUuLlapsed seuLlapeed seUuLlapeed seUuLlapeed seUuLIapeed seULIapeed seUuLlapeed seUuLIapaed seUuLIapsed seuLlapsed seULlIapsed seuLlapeed seUuLIapsed seuLlapeed seUuLIapsed seUuLlapeed seULlIapsed seUuLIapsed seUuLlapsed seUuLlapeed seUuLlIapaed seuLlapeed seUuLIapsed seUuLlapsed NO. 53, 2019 BANISTERIA 54 AAV ul SAV ‘HNWA U! HM9-VA dWM9-VA dWM9-VA HNWA Ul! Hdus-VA HNWA ul! Hdus-VA HM) u! Hdus-VA VA VA iVA iVA iVA GW aw GW GW GW GW Od 3d oe 8/8T ‘24U0D27 “1'[ SNLIOJUOD SNUILUE/Cd COST ‘a}U0De7 “TC sNDWed SN/IL{dould CQOQ8T ‘8]U0De7 “1'( 579ed0 Snjil{dould ZOST “JSIOYUSALI SAdHE/ SN/IL{GOUld (OST ‘uosysy) sediaid snuidosyje7 (8/8T ‘a]U0D27 “1'C) sN7e/aRI/IGO SsNIapaed QO8T “JSIOYUSsARID SNVe/ORN SNIADACd (€98T ‘d}U0D927 *1'C) sNUAUEedIAaS SdOIGOSSIT (SO6T ‘AdSe2) SNIILIBIIA SNSJEJOBELUOL (€98T ‘8]U0D27 “1'C) snUeLAUUId snsJeJOaeLUOY (So6T ‘AeseD) sdeaned snsJejOaewoy{ (ZO8T ‘JSJOYUdAeID) sodijed snsseJOIELWOHY (OP8T ‘UOSUIHF) SN/EYAaIOUR/AL SNSIEJOBELUOL (8/8T ‘a]U0D27 “T'C) SNUeEPLOY snsiejOaeWOH (8/8T ‘8]U0D27] “71'[) SILUOIINSY SNSJEJOBELUOHY (€98T ‘8}U0D27 “T'C) SNZE/GUO SNSJeEJOBCLWOHY (O€8t ‘AeS) SN79UID sNSJeJOBELIOL (OP8T ‘UOSYDIIF) SNUOIED SNSIEJOBELWOHY (sggt ‘Aases) ode) snsiejOaewioY (ZO8T ‘JSIOYUSACID) JO/OIIG SNSIEJOBELUOHY (ZO8T ‘JSIOYUSAPID) SNIpeG SNSs/eJOACLUOH 7SG8T ‘asyoes exopeled sISdodI[S (€9Q8T ‘A}U0D27 “7'[) SNSO.SUOL STUAPOULUE}S (9gT ‘a\U0De7 “T'C) SIP snjIbNYy L(68ZT ‘tINyAed) smENIIqUO SNYIENY (O8gT ‘2]U09e7 "71"C) sninzedo snbny TEST ‘Aes snjzequap snjibny L(6S8T ‘Zee41y) Ssisuaue/Aa2 snji6ny PULUIDO1d eul|lydould eul|lydould eul|lydould eul|lydould ePULIapeed PULIapeed euliqoydAlp euliqoydAlp euliqoydAip euliqoydAip euliqoydAip euliqoydAip euliqoydAlp euliqoydAp euliqoydAip euliqoydAlp euliqoydAip euliqoydAip euliqoydAip euliqoydAlp BUISdOdI|9S BUISCOdI|AS PUIDIIAS PUIDIIAS PUIDIIAS BUIDIIAS PUIDIINS julji4dould jul||4dould lulji4dould juljl4dould julji4dould |ULapeed |ULapeed |ULapeed |ULapeed |ULapeed |ULapeed ULapeed |ULapeed |ULapeed |ULapeed |ULapeed |ULapeed lULapeed |ULapeed |ULapeed |ULapeed 1UNIqO4Ue7 1UNIqO4Ue7 1UNIqO4Ue 7 1UNIGO4Ue7 1UNIqOJUe7 1UNIqO4Ye 7 1UNIqO4Ue7 seULIapeed seULIapeed seUuLlapeed seUuLlopeed seUuLIapsed seUuLIapsed seUuLIapsed seulIapsed seUulapsed seUulIapsed seUulIapsed seULIapsed seUuLIapsed seUuLIapsed seUulIapsed seUulIapsed seUlIapsed seUuLIapsed seUuLIepsed seUullapsed seUuLIopeed seULIapeed seULIopeed seULlapeed seULIopeed seULIapeed seUuLIopeed seULlapeed 55 BRATTAIN ET AL.: ROVE BEETLES OYo4 Ul! OSd-dW dWM9-VA HNWA ul! HM9-VA dWM9-VA ‘OSC ul 9sa-dW ‘NMA ul 9sa-0d WNSN ul 9sa-0d IOND UI 9Sd-VA OSC ul! OSCd-dW dWM9-VA AAV ul SAV ‘HM U! HM9-VWA VA VA idW GW GW 3d od od Od (Z68T ‘Aased) AILUYIS (SEPOLLIIGNd) SaposLeg (bbST ‘eaqny) suuNeYos (sapowayids) saposineg LhET Hed ‘OHaYaeyIs (Sapouiougeg) saposiyeg (6P8T ‘8]U0D27 “1'C) sdaauqgers (Saporerx7) saposimeg (pzgt ‘Aes) snuedu (sapouuougeg) saposiyeg (288T ‘Aasey) suoygound (sapouviougeg) saposiujeg (6P8T ‘a}U0D97 “1'[) sueoubiu (sapowiqnd) saposineg (CEST ‘eqny) syjoogeauy (SEepOAeIXZ) saposiyeg (6p8T ‘eqU0De7 “1'[) eeua/ (Sapo.nA/Z) Ssaposineg (O68T ‘japuetg) sme2Ny (SapOAeIXF) saposujeg (6p8T ‘a}U0De7 “1'L) SYE}UOY (SapOAeIXZ) saposineg (Z68T ‘Aase)) epnerissay (saponij2aq) saposiujeg (p8gt ‘AaseD) syjooquap (SapoUUuignd) saposiyeg (6p8T ‘eqU0De7 “1'C) SmeLASIG (SapOn[aq) saposieg Q06T ‘JayeeUurs *D sn7euUajUe (Sapouougeg) saposimeg (6P8T ‘8}U0D27 “1'C) SNSOASUOLU SAPOWILUASELYeg (6P8T ‘8]U0D27 “1'C) SIUYUOD sapolAseLyeg C68T ‘AQSED SNIN/OAU! SNILUYLIY 6b8T ‘8U0D2T “1°C SY/ODIGO/O snLULLIY OS6T ‘Wed ‘O yauueal sdoueLy LOP8T ‘UOSUDF SNELUOE SNUIAJOld ZG8T ‘asuoes snuequawe snuypebay/ (pPPST JAWIaUs|a, "J*4) Wwnuedvewe wrnjuobels (9gT ‘a}U0D27 “T'C) suUadiaid snydhyseyD ObST ‘UOSYydZ snazejsay snuNUe/ed eulsineg eulsiiyeg eulsiiyeg eulsiiyeg eulsiiyeg euIsineg eulsineg euIsLneg eulsineg eulsineg eulsineg euisiiyeg eulsiiyeg euisiiyeg eulsiiyeg eulsiiyeg eulsineg eulsineg eulsineg PULWIDO1d juisuyeg JuIsLyeg JuIsLyeg JuIsLyeg JuIsLyeg Juisieg juisuyeg juisueg juisuyeg Juisueg Juisuyeg JuIsLyeg JuIsLyeg JuIsLyeg JuIsLaeg JuIsLyeg juisueg luIsLyeg juIsLyeg luIdounewiy IUIU8}01q JUN2}O1d luj!4dould aeyisiyeg aeqsiyeg aeqsiieg aeqsiyeg aeqsiyeg aeyisueg aeqisuyeg aeyisiyeg aeyisiaeg aeyisiaeg aeyisiyeg aeqsiieg aeqsiieg aeqsiiyeg aeqsiyeg aeqsiyeg aeyisueg aeyisiyeg aeqisuyeg aeqisiaeg aeulydejasd aeulydejasd aeulydejasd aeulydejasd aeulydejasd aeulydejasd aeulydejasd aeulydejasd aeulydejasd aeulydejasd aeulydejasd aeulydejasd aeulydejasd aeulydejasd aeulydejasd aeulydejasd aeulydejasd aeulydejasd aeulydejasd aeulydejasd DEUIUID}OId DEUIUID}OId DEUIISIId SeULIEYIOIO|Ud seUuLlapeed NO. 53, 2019 BANISTERIA 56 HM) u! HM9-VA ‘HNA Ul! OSd-dW OAC u! OSd-dW OSC ul! ISA-WA dWM9-VA ‘OSC ul 9sd-dw VA VA idW aw GW GW GW GW GW Od od od 8/8T ‘2U0D97 “1'( S7eLRSIG (SISABDSOLIYY) SISAPISOUIYY (pest ‘Aase)) sayiuids snjzajdounkg (pest ‘Aasey) syenxas snjza/dounkd G/6T ‘AouBeN ‘“y'C pued snz2a/dounhkg 806T ‘Aesey syuuadibuo) snjzajdounkg (6P8T ‘8}U0D27 “1'C) suveauy sNnzdajdounKg (6P8T ‘23U0D27 “1'C) sIIdNwa}U sNIDajdOUuDKd L681 ‘Aasen snpapledu snjzajdounhd G/6T ‘souBem “yl s7e7/e/ snydajdounhg (6b8T ‘8}U0D27 “1'L) SYIOWYIP snDa/dounKq (pggTt ‘Aase)) sauUabuOD snjza/dounKd (pggt ‘Aese)) snuaziad snjza/d0}d97 p8st ‘Aasey syjoz/buo/ snz2a/dNnF (OTST ‘Yoequayoiey) Wuazsvey snyzajdnz (806T ‘Aased) SiLLoy sradejdnz L102 ‘UOWeN SapZebuaja snzIa/dNF 806T ‘Aesep “Np snz2a/dNnF 681 ‘Aasey snzountsip snjaajanz 6P8T ‘A}U0DeE7 “TL SsueNnyUoD snzoadnFz 806T ‘Aesep snuewioze snjzajdnz GOST ‘japuesg /A7U0Ia/ SOUCIPY 6P8T ‘83U0D27 “1'£ 57980) saueIpy (pest ‘Aesed) Se/UIBIIA (SAPOABIXZ) SAapOsLujeg (Z68T ‘Aase) sio2I0uN (sapouwougeg) saposijeg (O68T ‘lepuesg) seyynbueLy (SapoWignd) saposineg (Z68T ‘Aases) sy/elodiua} (saporeax7) saposiijeg (6b8T ‘a]U0D97 “1'L) SMeLyS (SapOWIgnd) saposLneg (6P8T ‘8]U0De7 “1'C) sNja/ds (sapouwougeg) saposijeg eulsdadsoulyy euuabiaelD eULaBIAe|D eulsiuyeg eulsineg eulsineg euIsLneg eulsineg eulsineg luIseldoja|\| juigDe]dN3 juqD9|dN3 juIqD9|dN3 juD9|dNy juQDe1dN3 juDe|dN3 juIq9|dn3 juDe|dN3 juIqD9|dN3 juyDe|dn3 juDe|dN3 juIyDe|dN3 juIqD9|dNn3 juQDe|dN3 juQDe|dn3 juIqD9|dNy juIqDe|dn3 juDe|dN3 juDe|dN3 lULaBIAe|D luUaBIAe|D juIsiyeg juisueg Juisuyeg juisuyeg juisuyeg juisuyeg aeypadny aeppadny aeypojdny aepoojdny aeypajdny aeppadny aeypajdny aeypajdny aeypadny aeypajdny aeypa|dny aepa|dny aeyoajdny aepoa\dny aeypajdny aeppajdny aeypajdny aeppajdny aeppajdny aeyoajdny ae WaBbie|D aeWwabieD aeyisiyeg aeyisueg aeyisuyeg aeyisueg aeyisueg aeqisieg aeulydejasd aeulydejasd aeulydejasd aeulydejasd aeulydejasd aeulydejasd aeulydejasd aeulydejasd aeulydejasd aeulydejasd aeulydejasd aeulydejasd aeulydejasd aeulydejasd aeulydejasd aeulydejasd aeulydejasd aeulydejasd aeulydejasd aeulydejasd aeulydejasd aeulydejasd aeulyudejasd aeulydejasd aeulydejasd aeulydejasd aeulydejasd aeulydejasd 57 BRATTAIN ET AL.: ROVE BEETLES HNWA Ul! HM9-VA OSC Ul OSd-dW HNWE U! ISA-VA ONI3 ul OSd-VA 9S ul 9Sd-VA SOVND Ul 9sa-dw TOND U! ISA-VA OSC ul! ISa-VA VA VA GW aw Od Od Od Od od od L(OT8T “YDequaysiey) ALWOIINAIG LUNILULL] (6p8T ‘a}U0De7 “1'C) E/qND eq/ao;LuLly (688T ‘Japuetg) E20e/0Y] eG/ay/ Z68T ‘Aasen enjezns eq/a (8791 ‘a4U0De7 “1'C) xe/duIs eqray (6ST ‘2]U0De7 “1'0) eed eqay L681 ‘Aase) sinuaz eyjasowujeg (896T “YOUeBUD ¥g 4a}SNUDS) /SaAGa7S SNUeSOLL/EG (S98T ‘lepueig) LwnjzezUNdUy LuNhoY (€96T SHed ‘O) Heubem sajsenoy (6b8T ‘a1U0De7 “1'C) 4ayigo/b sajsenoy (8Z8T ‘8}U0D97 “1'L) SY/ODIANQY SaRSeHOY 068T ‘jepuesg sMpa/oSgo snjDe/GoNUL] 6b6T ed ‘O/YIeG/ane sNyDa/AO/wLl | (9gT ‘2]U0987 “1'C) sUuOyINe? WNISAL/] (O68T ‘Japuatg) sN7e;NSSOY SAISEISAYL (S98T ‘Japueig) eyULO eDaweYy (€68T ‘japussg) LuNaVOgGeled winoepnasd (6p8T ‘8}U0D27 “7'C) wnjenae wng7zepnasd 9G6T “4d ‘O SNDDeI0Y] SsN/YAAINT O88T ‘a}U0De7 “TL SY/LUIs snjYaAINF POET ‘AeyeY AULUYOS srjyYaAAINZ €68T ‘AaseD suauiuosd snjyarjn7 (ZS8T ‘Aysinyds}Op|) sdeayr (sapojajdojqig) snyoajdojqug (pest ‘Aase)) sdeaina/ (sapojajdojqig) snzoajdojqrg (8/8T ‘a]U0D27 *1'f) 4ebajzl (sapojpajdojqug) snjoadoyjqig 066T ‘Ja|pueUD Mez0YI snzDa/doO/qQIg PUIILUL | PUIILUL | PUIILUL | PUIILUL | PUIILUL | PUIILUL | PUIILUL | PUIILUL | PUIILUL | PUIILUL | BUIILUL | EUIILUL | eunueydeued eunueydeued eunueydeued eunueydeued eunueydeued eunueydeued eunueydeued eunueydeued eunueydeued eunueydeued eunueydeued eunueydeued eunueydeued eunueydeued eunueydeued IUIYDAUOUSL | IUIYDAUOYSL | IUIYDAUOUSL | IUIYDAUOYSL | IUIYDAUOYOL | IUIYDAUOYSL | IUIYDAUOYSL | IUIYDAUOYSL | IUIYDAUOUSL | IUIYDAUOYSL | IUIYDAUOUSL | IUIYDAUOYSL | IUIYDAUOUSL | IUIYDAUOYSL | IUIYDAUOUSL | IUIYDAUOYSL | lUIYDAUOYSL | lUIYDAUOYSL | IUIYDAUOYSL | lUIYDAUOYOL | IUIYDAUOYSL | IUIYDAUOYSL | IUIYDAUOYSL | IUIYDAUOYSL | IUIYDAUOYSL | lUIYDAUOYSL | lUIYDAUOYSL | aeypajdny aeypajdny aeypajdny aeppadny aeppoadny aeypadny aepoa|dny aeypodny aeyooajdny aeypajdny aeyoa|dny aeypajdny aeyo|dny aeyypadny aeypajdny aeypadny aeypajdny aeypadny aeyoajdny aeyoadny aeypoajdny aeppadny aeyoajdny aeypadny aeyoojdny aeypajdny aeyoojdny aeulydejasd aeulydejasd aeulydejasd aeulydejasd aeulydejasd aeulydejasd aeulydejasd aeulydejasd aeulydejasd aeulydejasd aeulydejasd aeulydejasd aeulydejasd aeulydejasd aeulydejasd aeulydejasd aeulydejasd aeulydejasd aeulydejasd aeulydejasd aeulydejasd aeulydejasd aeulydejasd aeulydejasd aeulydejasd aeulydejasd aeulydejasd NO. 53, 2019 BANISTERIA 58 OZOW Ul! 9Sd-9d HNWA Ul! OSA-WA HM) Ul SIW-VA OZOW Ul! OSA-WA VA VA iVA GW dW GW oe Od Od 3d (€68T ‘jepueig) evespUy/AD e1DEqUEYIAY (S9O8T ‘Japuetg) AUABUOD eIYIEQUAYIIAY L681 ‘Aase) sealing elljpequayaey (O68T ‘Japuaig) 1E/0I1g BIYDEQUEYIIIY (998T ‘jepueig) Baquege enjpequeyriey (EBT ‘aqny) esojUaLwo} SIxesi\y GTOz ‘epond wweyyom (en/bAyseig) eyn/bAyoesg (998T ‘jepuaig) sayin (epn/bAyseg) eyn/bAyerg (c6gt ‘Aase) epeigaiay (eyn/bAyoesg) eynjbAyoeig G10Z ‘Ja|pueyD aeumeys (eynj/bAyoesg) eynjbAyoeig (6p8T ‘eqU0De7 “1°0) abun] (esiyy) enjbAyse1g (998T ‘japusig) eypeuvazu (eyn/bAyoe1g) enjbAyoesg (S9O8T ‘japussg) eEuepLUOY (eyn/bAyoelg) eynjbAyoesg (pzgt ‘Aes) eyequap (eqnbAyoeig) eqnjbAyoelg (ZS8T ‘AYS|NYISJOP|) SLLVIAA/LUOD (eyn/bAyoesg) eynjbAyoeig (G98T ‘lepuesg) syOIINeD (esi\y) eN/bAYelg (O8gT ‘eqU0De7 “1'f) abeyjaq (emnjbAyoelg) enjbAyoeig (2681 ‘Aasep) enbue (eqn bAyoeig) eqnibAyoesg (CEST ‘eqny) syeunuopge (eynjbAyoeig) eynjbAyoeig OTOZ ‘UOIIED 1B O94 /UYYIAYS] PLUOUOS OTOZ ‘UO ED 9 Oa SN//EYAOIOYAOJAANS PLUOUOS OTOZ “UOIIED 8 OLUDY /SOLUOY PLOUOS (Z68T ‘japuetg) saplosajsebo.y snixayseoqny (ZEBT ‘JUDI9I4) SNZOSVAd SNIxayseOqnZ (6P8T ‘8}U0D27 “1'C) smje/nayeue? snjda/douo7 SLET ‘83U0D27 “T'[ SELISGNS SNIXOYY E68T ‘|OPUAIG MA/LUYIS SNIXBYY 6b8T ‘84U0De7 “TC SN7a/NISUI snIxEeyy 806T ‘Aasep snau/bn.uey snIxayYy euln|bAuseig euln|bAuseig euln|bAuseig eulni|bAuseig euln|bAuseig euln|bAuseig euln|bAuseig euln|bAuseig euln|bAuseig euln|bAuseig eulnj|bAuseig euln|bAuseig euln|bAuseig euln|bAuseig euln|bAuseig euln|bAuseig euln|bAuseig euln|bAuseig euln|bAuseig eulsebo | eulnsebo | euLsebo | BUIIXOUY eUIIXeyy eUlIXeyy eUIIXOyy lunn|bAyse1g lunn|bAyse1g lunn|bAyseig lunn|bAyseig lunn|bAyserg lunn|bAyse1g lunn|bAyse1g lunn|bAyse1g lunn|bAyseig lunn|bAyse1g lunn|bAyse1g lunNn|bAyse1g lunn|bAyse1g lunn|bAyse1g lunn|bAyseg lunn|bAysesg lunn|bAyse1g lunn|bAyse1g lunn|bAyse1g juLSeBOt | iuULNSeBO | 1Ul4}SeH04 | iuLISeBOt | 1Ul4}SeHO4 | juLSeBOt | 1ul4}SeHO4 | deILBNeIUOD de1e0eIU05 de1B0e1U05 de1B0eIUOD deqe0eIU0D de1B0eIUOD de1B0eIUOD deqa0eIUOD deILBNeIUOD de1B0eIU0D deIBNeIUOD deIBOeIUOD de1a0eIU0D de1BNeIUOD de]B0eIU0D de1La0eIU0D deIBNeIUOD de1B0eIU0D de1a0eIU0D aeyuose4 aeyuose4 aeyuose4 aeypojdny aeypadny aeypadny aeyoa|dny aeppajdny aeyoodny aeypajdny aeulydejasd aeulydejasd aeulydejasd aeulydejasd aeulydejasd aeulydejasd aeulydejasd aeulydejasd aeulydejasd aeulydejasd aeulydejasd aeulydejasd aeulydejasd aeulydejasd aeulydejasd aeulydejasd aeulydejasd aeulydejasd aeulydejasd aeulydejasd aeulydejasd aeulydejasd aeulydejasd aeulydejasd aeulydejasd aeulydejasd aeulydejasd aeulydejasd aeulydejasd 59 BRATTAIN ET AL.: ROVE BEETLES HNWA ul! OSa-VA OSC ul ISA-VA OSC ul! ISa-WA OSC ul! ISA-VA HNWA U! HM9-VA OAL u! OSd-dW OZOW U! OSA-VA OSC ul ISA-VA OSC ul! OSCd-dW HNWA Ul! OSa-VA OZOW ul! 9Sd-9d VA VA GW GW od od od (6b8T ‘2]U0D87 “T'[) unU_Njan UoJYLIEIEg GOST ‘JapUatg LUNSOLUBAS UOIY2IEIIG (68 ’2]U09e7 “7'1) wnynbuoy volypezag SG6I Hed ‘OWwnjsnl uolypeIeg QG6T “Wed ‘O /uapmoy uolypyie7aq (6pP8T 12]U0D07 “1'C) PaqLLUOY UOIYjIEIAg GOST ‘Jepueig LWNpaSxa UOIYLeIEG (6P8T ‘2}U0D27 “T'f) euuwouge UosYyzIeEIEGq (O68T ‘JapUatg) S/LIODLIEA sIxegAy (O68T ‘jepussg) EDI/EA sixeghy L76T ‘||24 esvansuey sixeqry LEI “|1e4 suapanbigo sixeqrhy (68 ‘a]U0De7 “1'C) euNfuo2 sixeghy (S98T ‘jeapuasg) Bere sixeghy (prst ‘eqny) epunougns e1zequeayrey (O88T ‘e}U0De7 “T'L) suepes ej2equayIeYy (6P8T ‘2]U0987 “1'C) syoomound elysequayay (6p8T ‘a}U0De7 “1'C) enbuidoid enljsequayrleYy 1681 ‘AaseysojeuLibBalad e1IeGUAYWIEY £007 ‘UO WeD eUeNsINO] e1yDeqUEeYIIAY L68T ‘Aase euesuey eIYDEqUaYIIEY (pest ‘Aase) edo eIyDeqUAaYIIOY QG6T ‘4ed ‘OJPIEMOY eIYDEQUAYIIOY OG6T Hed ‘OMprey elyo2equeyrey (pest ‘Aese)) syoe/b enljzequayriey (pest ‘Aases) sie eIYDEQUAYIIAY (O88T ‘a}U0De7 “1"[) suabranip eIyoequayrlay C68T ‘Aasen essiuap elyDequayolay euLyedEq euLyLe.Eq euLyLe.Eq euLyLe.Eq euLyLe.Eq euLyLe.Eq euLyLe.Eq eulype.eq euln|bAuseig euln|bAuseig euln|bAuseig euln|bAuseig euln|bAuseig euln|bAuseig euln|bAuseig euln|bAuseig euln|bAuseig euln|bAuseig euln|bAuseig eulnj|bAuseig euln|bAuseig euln|bAuseig euln|bAuseig euln|bAuseig euln|bAuseig euln|bAuseig eulnj|bAuseig euln|bAuseig lunn|bAyse1g lunn|bAyse1g lunn|bAyseig lunn|bAyse1g lunn|bAyse1g lunn|bAyse1g lunn|bAyserg lunn|bAyse1g lunn|bAyseig lunn|bAyseig lunn|bAyserg lunn|bAyseg lunn|bAyse1g lunn|bAyseg lunn|bAyse1g lunn|bAyse1g lunn|bAyse1g lunn|bAyse1g lunn|bAyse1g lunn|bAyseag lunn|bAyse1g lunn|bAyse1g lunn|bAyse1g lunn|bAyse1g lunn|bAyserg lunn|bAyse1g lunn|bAyse1g lunn|bAyse1g de1B0eIU0D de1BNeIUOD de1a0eIU0D de1BNeIUOD de1IB0eIUOD de1B0eIUOD de1a0eIUOD de1BNeIUOD deqLa0e1U05 deqWa0eIU0D de1a0eIU05 de1B0eIUOD deqa0eIU0D de1BIeIUOD de]B0eIUOD de1LBDeIUOD de]a0eIU0D deqa0eIU0D deqa0eIUOD de1B9eIUOD de1LIB0eIUOD de1a9eIU0D deIBNeIUOD de1B0eIU0D de1LBNeIU0D de1B0eIUOD de1a0eIU0D de1a0eIU0D aeulydejasd aeulydejasd aeulydejasd aeulydejasd aeulydejasd aeulydejasd aeulydejasd aeulydejasd aeulydejasd aeulydejasd aeulydejasd aeulydejasd aeulydejasd aeulydejasd aeulydejasd aeulydejasd aeulydejasd aeulydejasd aeulydejasd aeulydejasd aeulydejasd aeulydejasd aeulydejasd aeulydejasd aeulydejasd aeulydejasd aeulydejasd aeulydejasd NO. 53, 2019 BANISTERIA 60 HNWA ul! HM)-VA ONIS u! OSA-VA HNWA Ul! ISA-VA OSC ul! OSa-WA OSC ul! OSd-dW OSC ul! OSA-VA iVA iVA GW Od od od (258 ‘Aysinusjo;) sap/oisLgeg eunjodn ZEBT ‘JeUdIa|4 SNLUIdO (SapIOJOWeLH) sNjJOweLH 6P8T ‘8U0DeET “TL SW/e}SOD SNIOYAISAWW J (p7gt ‘A@S) SNICEULIED SNUOYAISOLU | 6P8T ‘A4U0De7 “TC /uosYyauE snydejasd C6gt ‘Aasey xeyag snydejasd (6P8T ‘8}U0D27 “1'L) /UELLUALULUIZ SAapOSIUaZD (6P8T ‘e]U0De7 “1'[) snazd saposjuazD (4681 ‘Aase)) SLISNIe/ SapOsiUazD (6P8T ‘8}U0D927 “1'L) SNULIGOSUOD SaposiuazD (998T ‘JapUetg) S/LODI/IUOLU SnNUAY (POET “AesYeu) SYELUAIS SNYDARHIICAN (pest ‘Aesea) snazejsa} sNyaAjOyony] 886T UalpueUD syejUALIO SNIDIeIPUI|AD (6p8T ‘a]U0De7 “1'L) siaediBuo) snj2sespuyAD €68T ‘Aasen /AV/ULO SnjzsesPUI|AD (€68T ‘Adse)) SYEILIAA SNYDAJO}SND (Z68T ‘Aasez) sequoyerod snydAjojsnz OTOZ ‘IUIWO1a5 9g Ja|pueYD ZaYyI7eU SNYDAJOISND (6p8T ‘ayU0De7 “7'C) JOU SNYDAIOISND OTT ‘IUIWOLa5 1g JajpueUD s/ego/ SNYDAJO}SND (661 Hed 'O) Ab6ep snyshjojsnZ 9T0Z “IUIWOJa5 1g Ja|PUeYD LWoeoped snyrAjojsNnZ (Z68T ‘Aasep) aeuyjoser snuiyyAGOYyoA 1 (S9O8T ‘Japuetg) sNJeULED SapOIBeYyIeEy (S98T ‘japuesg) SNPeEUOZ SAZYIOLVINT ZS8T ‘Aysinyosjop| s7ejeyp sniuasdnz PUIUOICIJELUOS PUIUOICI}ELUOS 1ULA | 1ULA | lULOYCISSW | lULOYCISSUW | lulydejasd lulydejasd lUIISIUSID JUNSIUDID [UIJSIUBYD JUIISIUYD JUIISIUSYD IWIYOA | IUIYDA | IUIYOA | IUIYDA | IUIYDA | IWIYOA | IUIYOA | IUIYDA | IWIYOA | IWIYOA | IUIYDA | IUIYDA | lUIUIyyAg lUIUIyAg lunn|bAyserg lunn|bAyse1g aeyydejasg aeyydejasd aeyydejasd aeyydejasg aeyudejasd aeyydejasg aeyydejasg aeyydejasg aeyydejasd aeyydejasg aeyydejasg 9EWaDeIUOD dE WIBDeIUOD VE WVIeIUOS) Je BIeIUOS) JE WIVIeIUOS) JE IBVIeEIUOS) JE WIBIEIUOS) VEWIBIEIUOS) Je WIIeIUOS) VE WIBIEIUOS) JE WIBIeIUOS) JE WIBIEIUOS) JEWIVIeIUOS) JE WIBIeIUOS) JE WIVIeIUOS) JE WIVIeIUOS) aeulydejasd aeulydejasd aeulydejasd aeulydejasd aeulydejasd aeulydejasd aeulydejasd aeulydejasd aeulydejasd aeulydejasd aeulydejasd aeulydejasd aeulydejasd aeulydejasd aeulydejasd aeulydejasd aeulydejasd aeulydejasd aeulydejasd aeulydejasd aeulydejasd aeulydejasd aeulydejasd aeulydejasd aeulydejasd aeulydejasd aeulydejasd 61 BRATTAIN ET AL.: ROVE BEETLES dWM9-VA dWM9-VA dWM9-VA HNWA Ul! ISA-VA VA iVA iVA iVA aw GW GW od Od 3d (Z68T ‘Aese)) SNILIBIA SApOULIaYAga7Z (ZS8T ‘23U0De7 “1'C) sNSo/od0D sapouuaydaz O98T ‘8}U0D27 “TL Sap/oseumueB WnIpIxo | PrST ‘JaWIaYs|ap| "Q*4 WwnpeUNLUA, eELUOSIYdeIS 098T ‘a}U0De7 “TL ae/nzns ewosiyders O66T ‘1407 8 UBYDSE7 /WeYdaps eLUOSIYAeIS O98T ‘8}U0D27 “TC wnysnd ewosiyders O98T ‘8}U0D97 “TC wnjenjound ewosiyders GZ8T ‘AS LUNXAAUIOD BLLIOSILY{AeIS (Z96T ‘||JQUIOD) eUWayINIAsS e1az0aeg (Z96T ‘||QUIOD) e7/ADS e1a200eg C68T ‘AaseD earid e12a908eg C68T ‘Aasep eveu e1az0aeg E66T ‘UeYdays 1g [GQ] BUASIPUY Bla00aeg OZ6T ‘pseuoy e7esyey e1az0aeg C68T ‘Aasea exayap elaz0aeg C68T ‘Aaseyvauebuo) e1az0aeg (L96T ‘||QUIOD) Epgtieyo e1a200eg 098T ‘aqU0De7 “TL sve2Z/de e1a00aeg 006T ‘Aase> syeumuopge e1a008eg EzeT ‘Aes winjeynbupenb wnipiyders bpST ‘Jawlaysja,| "y*4 wed wripiyders (TOST ‘snidluqe4) sojoau0D LunuiedAD G/6T ‘URWWIAH e7e/ngns sisdopnasd Z68T ‘Aesey daqgniUuas SrA (bST ‘aqny) sye/aluny snsh 1 6b8T ‘a}U0D27 “TC sy/uoW snyjAydoaz 6b8T ‘aqUODe7 “1"C ve/balz (sNIpaDOASIUIS) SNIpAD 6b8T ‘a}U0De7 “T'[ sNsouds (snipaD) snipap PULAL ePULAL PULA, PULA, ePULIAL lulluuaydes lulluusydes luUeWOSIYdedS luNeWIOSIUdeIS luNeWIOSIYdedIS lueWIOSIUdeIS lUNeWOSIYdeIS luNeWIOSIYdedS luNeWOSIYdedS lueWOSIUdedS lueWOSIYdeIS lueWOSIUdedS luNeWOSIYdedS lueWOSIYdedS lUeWIOSIYdedS luUeWOSIYdedIS lueWIOSIYdedIS lueWOSIYdeIS lueWOSIYdedS luUNeWIOSIYdeIS lulIplydeos lulIplydeos juiuedAy IAL IULA | IAL IAL IAL aewuuaydey aeWuUsydey aeyydejasg aeyydejasg aeyydejasd aeyydejasd seyydejasd aeulusewpAds aeulusewpAds eeullplydess eeullplydess eeuliplydess eeullplydess eeuliplydess eeuliplydess eeuliplydess eeullplydess eeuliplydess eeullplydess seullplydess eeullplydess eeullpiydess eeulipiydess eeullpiydess eeulipiydess eeuliplydess eeullplydess eeuliplydess eeuliplydess eeuliplydess aeulsdopnasd aeulydejasd aeulydejasd aeulydejasd aeulydejasd aeulydejasd NO. 53, 2019 BANISTERIA 62 VA GW GW Od Od Od L681 ‘Aases sinua} snipeleaN (ZS8T ‘83U0D27 “1'L) SN//ESILU SNLUPAISOIN 866T ‘2J22y,0 ase? SNLUPADSOLOIP/ GO6T ‘aeeyns “D sNnaqUeye srapolydo7 681 ‘Aasey syemuan (snuuoanz) snuuognz (Z68T ‘Aases) sayuLy (snyzodeny) snuuoanz (Z7S8T ‘a}U0D927 “1'L) so7eu/es (snydowiosg) snuuognz (ZS8T ‘aqU0De7 “1'C) snse/ (snYdoudDAg) snuuognz (€98T ‘a]U0De7 “1'C) syepiuesd (snyz0dey) snuuoanz 681 ‘Aasey $17/N220 (snuuoINFZ) snuuognz (ZS8T ‘a}U0De7 “T'L) sNy/ainasgo (snysodey/) snuuodInz (998T ‘ssngneyos *\"]) /a7zuaDa/ (SNUUOINFZ) snuuOoINF (2681 ‘Aase)) snnzouul (snyzodenxy) snuuoonz (Z68T ‘Aases) syiuny (snysodey|) snuuoonz (ZS8T ‘aqU0De7 “1'C) sNyeLY (snyzoden) snuuoagnz (ZS8T ‘a]U0D97 “1'C) snajry (snYodenxy) snuuO DNF (ZS8T ‘a]U0D27 “1'C)4abissoy (sny20dey/) snuuoINz (ZS8T ‘8]U0D27 “1'C) sisveyAey (snyzodeny) snuuoanz (ZS8T ‘aqU0De7 “1'C) sANjey (snydowosg) snuuodnz (ZS8T ‘8}U0D27 “1'L) sMULgosUOD (snuuoINZ) snuuOINF (pzgt ‘Aes) sadiaea (snuuognz) snuuoonz (ZS8T ‘aqU0De7 “1'C) smjeAe— (snyz0den) snuuoanz (ZS8T ‘a]U0D97 “1'C) sn/nsoyideo (snyz0den) snuuoanz (Z68T ‘Aase)) snyesezes (snyzodey|) snuuoonz (pzgt ‘AeS) syoainaig (snyoodeny) snuuoonz (ZS8T ‘a]U0De7 1'C) syeseg (snyz0dey/) snuuognz (ZS8T ‘aqU0De7 “1'C) syeue (snyz0denxy) snuuooNz Z68T ‘Aasea sninjsngos snijag (ZS8T ‘aqU0De7 “1'C) e7ez2UNAdQns sisdaaAyoe/g 998T ‘A}U0D27 “1'[ eEUaOWe ee/OINIYD (6ST ‘jepuesg) sN7e/ snUODIYIN lule|npue|s lule|Npue|S) lule|npue|S luje|Npue|S) lule|NpUue|S lule|Npue|S) lule|NpUue|S) lule|Npue|S) lule|Npue|s lule|Npue|s) luIe|NpUue|S) lule|Npue|s) lule|Npue|S) lule|Npue|s) lule|Npue|S lule|Npue|S) lule|Npue|S lule|Npue|S) luje|Npue|s) lule|Npue|s) lule|Npue|s) lule|Npue|S) luje|Npue|s lule|Npue|S) lule|Npue|S) luIe|NpUue|S) luje|Npue|s lule|Npue|s) luIe|Npue|S) 1UIE|OAABYD juayyny JEWUBELUPAIS BJeWUBELUPAIS SJEWUBELUPAIS JeEPWUSELUPAIS SJeWUBELUPAIS SJEWUBELUPAIS SEPUBELUPAIS SJEWUBELUPAIS SeWUBELUPAIS JePWUBELUPAIS SJeEPUBELUPAIS JeWUBELUPAIS SeWUBELUPAIS SJePWUBELUPAIS SeEPWUBELUPAIS SePWUBELUPAIS SJEWUBELUPAIS JeWUSELUPAIS SJEWUBELUPAIS SeEPWUBELUPAIS SJEPUBELUPAIS JeWUBELUPAIS JeEWUBELUPAIS DEYWUSELUPAIS SePWUBELUPAIS SJeWUBELUPAIS SEYWUBELUPAIS SJEPWUBELUPAIS SJEPWUBELUPAIS JeEWUBELUPAIS aewuuUsydey aeurusewpAds aeurusewpAds aeulusewpAds aeurusewpAds aeurusewpAds aeurusewpAdsS aeulusewpAds aeurusewpAds aeurusewpAds aeurusewpAds aeurusewpAds aeulusewpAds aeurusewpAds aeurusewpAds aeurusewpAds aeulusewpAds aeulusewpAdS aeulusewpAdsS aeurusewpAds aeulusewpAds aeulusewpAds aeurusewpAds aeulusewpAds aeurusewpAdS aeulusewpAds aeulusewpAdS aeulusewpAds aeulusewpAds aeurusewpAds aeulusewpAds aeurusewpAds 63 BRATTAIN ET AL.: ROVE BEETLES HNWV ul NAV-VA dWM9-VA HM) Uu! HM9-VWA dWM9-VA dWM9-VA VA VA VA VA aw GW GW GW GW GW od 3d Od od od (TO8T ‘Sninuge4) siuuadyns snyonuojag GL6T ‘YURI "H'C SNOMSN SNILVOSYOLI G/6T ‘UCI “H'L SN//ASO/ SNIVOSYDUF (P88T ‘UIOH) OsNd sniUOSYOUZ (PRgT ‘WOY) eyazed sniuOsyQUZ (PSST ‘WIOH) sn2/ed SNIVOSYDUF (P88T ‘UIOH) SNUEU SNIUOSYOUFT GZ6T ‘YUP “H'L SNDIAID SNIUOSYIUT G/6T “YURI “HL S/CYA2DAYIEIG SNIUOSYQUT G/6T ‘Ue "H'C SNULUNe SNIUOSYDF (OP8T “UOsUdIIZ) syjoaWoUNd snuoydouedus | CQ8T ‘9]U0D27 *T'[ O/sNd sdoYyjO1EIA} ZT0Z ‘eue}aWs 9 ayUNIg SsNZUYLI SnipanbiLuaH (8/81 ‘a]U0De7 “1'[) X/iWeA snipanbeuy OPST ‘uosydy snuaid snioydojAIy CST “JBIAIANG /azUO0Ig/ srUOYdo/ADy ZS8T ‘asudeS SI//ODIAEY SrUOYAO/ADy 8/8T ‘a}U0D27 “T'( snsuep snuoydoj~Aoy OZ6T ‘bua] ases snioydojpoy TZ6T ‘euejaws syibe snioydojAoy (906T ‘AeseD) snuequeWwe snoe/y 09ST ‘Zeesy MLUNeYIS SNYD0IG Tp8T ‘WINEUIS /UUELLUALULUIZ SNUBELUPADS (€9gT ‘a]U0De7 “T'L) YSINYOSJOW SNUBELUPAIS (Z68T ‘Aese)) syjonbuo/ snuaeupArrs (€98T ‘A}U0DE7 “1'L) SNSSOIB SNUBELUIPAIS L68T ‘Aase) SUOYIAED SNLUPADSO}AIT (TP8T ‘wneuds) smje/opiad SNUYIIUAIS (ZS8T ‘8]U0D927 “1'0) snue/qua snuyoIUuays (Z68T ‘Aasey) snipeg snuyoiuays PUIYJUO}IUd PUIIUOSYDIIF PUIIUOSUDIIIF PUIIUOSYDIIF PUIIUOSYDIIF PUIIUOSUDIIF PUIIUOSUDIIF PUIIUOSUDIIF PUIIUOSUDIIF PUIUOSUDF PUIUIJOSIUY eululdoA|quiy euuoydojAoy euuoydojAoy euuoydojAoy euuoydojAoy euuoydojAoy euuoydojAoy euuoydojAoy euuoydojAoy julul|Aydeys julul|Aydeys juluI|AYydeys julul|Aydeys julul|AYydeys julul|Aydeys julul|Aydeys julul|Aydeys julul|Aydeys julul|Aydeys juluI|AYydeys juluI|AYydeys julul|Aydeys julul|Aydeys juluI|AYydeys julul|AYydeys julul|AYdeys julul|Aydeys julul|Aydeys juluI|AYydeys [UIIYIO lUIYooIq lUIUaeLUPADS |UIUBeLUPADS lUIUaeLUPADS |UIUBeLUPADS lulLupAdSo}de7 luLe;npue|s) julLe;npue|s) juILe;npue|s) SeEWUBELUPAIS SEPWUBELUPAIS JeWUSELUPAIS JEPWUBELUPAIS SJEWUBELUPAIS SJeEWUBELUPAIS JEWUBELUPAIS DJEYWUSELUPAIS aeurulAydeis aeuruljAydeys aeuruljAydeys aeuruljAydeys aeuruljAudeis aeuruljAydejys aeuruljAyudeys aeuruljAydejys aeuruljAydeys aeuruljAydeys aeutuljAydejys aeuruljAydeys aeurulAydeys aeutuljAydeys aeuruljAudeys aeuruljAydeys aeurulAydeys aeuruljAydejys aeuruljAydeys aeuruljAydeys aeutuljAydejys aeutuljAydeys aeurusewpAds aeurusewpAds aeurusewpAds aeurusewpAds aeulusewpAds aeulusewpAdS aeurusewpAds aeurusewpAdsS NO. 53, 2019 BANISTERIA 64 dWM9-VA dWM9-VA dWMD U! Hdus-VA iVA VA GW GW GW dW GW GW od od 3d Od od od (O€8T ‘WiayiouUeLp|) SUUada/NiaeD SNYJUO/IYd P88T ‘UIOH 4adse SNYILO/IYd P88T ‘WOH syvenbae snyzUoliYd (€98T ‘28]U0D27 “1'[) SYEUILLLIA] SNIUSIGOaNY (OP8T ‘UOSYDIIZ) SMULIGOS snIUsIgoayy (€9gT ‘a}U0De7 “1'[) Sapiasapaed sniusiqoay TS6T ‘yUeI4 “H'[ Sap/ojUapInI0 sniusigoayy (€9QT ‘a]U0De7 “1'C) Sr/Npides sriusigoaN (p88T ‘UIOH) snpunont sniusigoaN (P88T ‘UWIOH) sNsoz0l sniusiqoaN (OE8T ‘AeS) snynjoe/ snyguOjnjae7 SS6T ‘a100W WYyajs sruadsoy (ZO8T ‘JSIOYUSACID) SisUaIOLUIV/EG Sredsay] (OEgt ‘Aes) syenide sniedsayy 4(OS8T ‘eqny) wnieuuay] snyjuosqey G66T “EURJOLUS SNIGIA SNLIGe L(ZO8T ‘JsyOUUsAeID) SN/NpIpUa/ds snugey (O€6T ‘Ile4) sdeoyero snugey A(ZO8T ‘JSsHOyUdAeID) sN/NqbiU snugey (p88T ‘WOH) snujeYyzydoinU snuige9 (P88T “WJOH) snsoneyey snuge5 LZP6T ‘ZJeduaeurs snzeLySNe snugey (OST ‘UOsYysHA) sNjzeLASIG sNIeLYSIG SNYeD I(ZO8T ‘JSIOYUSACIN) SNPIDIOS SNIUSIG (€pST “‘WiaysouUeW)) Wp/embals sniusig (ZS6T ‘YdIeH) sisuajabnd sniusig (SS6T ‘eueaLUS) LL/ed sniusig (Ob8T ‘UosYyoy) snzainbul sniusig I(ZO8T ‘JSsoyUdsAeID) sazo/eydan sniusig (908T ‘JSu0YUaAeID) sNpUe/G sNIUsIg PUILUO|Id PUILUO|IUd PUILUO|IUd PUILUO|IUd PUILUO|IUd PUILUO|IUd PUILUO|IUd PUILUO|IUd PUILUO|IUd eUILUO}!Ud PUILUO|IUd PUILUO|IUd PUILUO|IUd PUILUO|IUd PUILUO|IUd PUILUO|IUd PUILUO|IUd PUILUO|IUd PUILUO|Id PUILUO|IUd PUILUO|IUd PUILUO|IUd PUIUUO|IUd PUILUO|IUd PUILUO|IUd PUILUO|IUd PUILUO|IUd PUILUO|IUd PUILUO|IUd PUILUO|IUd lululAydeys julul|Aydeys lulul|Aydeys lulul|Aydeys julul|AYydeys lulul|Aydeys lujul|Aydeys julul|Aydeys juluI|AYdeys lujul|Aydeys lululjAydeys julul|Aydeys juluI|Aydeys lulul|Aydeys lulul|Aydeys juluI|AYydeys julul|Aydeys lulul|Aydeys lululAydeys julul|Aydeys julul|Aydeys lulul|Aydeys juluI|AYydeys julul|AYydeys lulul|Aydeys lujul|Aydeys julul|AYydeys lujul|Aydeys lulul|Aydeys julul|Aydeys aeuruljAydej}s aeuruljAydej}s aeuruljAydej}s aeutuljAydejys aeuruljAyde}s aeuruljAydej}s aeutulAydeys aeuruljAydej}s aeuruljAydej}s aeuruljAydej}s aeuruljAydej}s aeuruljAydej}s aeuruljAydej}s aeuruljAydej}s aeuruljAydejs aeuruljAydej}s aeuruljAydej}s aeuruljAydejys aeuruljAydej}s aeuruljAydej}s aeuruljAyde}s aeuruljAydeys aeuruljAydej}s aeuruljAydej}s aeutuljAydeys aeuruljAydej}s aeuruljAydej}s aeuruljAydej}s aeuruljAydej}s aeuruljAydej}s 65 BRATTAIN ET AL.: ROVE BEETLES dWM9-VA dWM9-VA VA VA aw aw GW oe od Od GI6T ‘AaSeD SNpI/EA SNYJUO/IYd (ZO8T “JSIOYUSACIS) SNULIQLUN SNYJUO/IYd G66T ‘CURJLUS SAP/OUILIGLUN SNYJUOIIYd L(ZO8T ‘JSIOYUSACI) SYMEIQLUN SNYZUO[Yd (ZO8T “JSIOYUSACIN)) SNDICIOY] SNYJUO//Yd P88T ‘WOH SNUIDUAS SNYIUO/IYd (ZO8T ‘“JSIOYUSAPID) SUEIVIAS SNYJUO//Yd PST ‘UWIOH SN/NYN SNYJUOLIYd IpZ8T ‘daeys snjnbuezas SNyYJUOl/Yd P88T ‘UWIOH sy/ooupenb snyzuojild L(8GZT ‘snaeuuly) smqjod snyzUojiUd (908T ‘JsJoyUaAesD) snzeY/ed SNYUO/IYd GOGT ‘CURJBLUS SIJEUOBU SNYJUO/IYd G66T ‘CURJALUS SASBELOLL SNYJUO/IYd LZERT ‘Suaydays S/LWODBUO/ SNYJUO/IYd OP8T ‘UOSUDHF SIELUIO/ SNYJUOIIYd G66T ‘URS SUL SNYJUO/IYd OPST ‘UOSUDHA sNaVedoy SNYJUO/IYd GT6T ‘Aaese) voyi9e/B SsnyJUoIYd G66T ‘CURJBWUS SMAED SNLQUO/IYd PBT ‘JOWIOYS|AW “A'4 SLLUOJISN SNYJUO{Yd GI6T ‘AdSe> SNEUILUNY SNYJUO/IYd GI6T ‘AdSe> SISEGIACY SNYJUOIIYd I(ZO8T ‘JSIOYUSAPIO) SNAPIOISID SNYJLOLIYd L(ZO8T ‘JSIOYUBARID) SIIGAP SY JUOLYd P88T ‘WOH SUEJIUND SNYQUO/IYd L(O6ZT ‘UlJAWID) S77e7UANIO SNYJUO/IYd I(ZO8T ‘JSIOYUSALIO) SNUUIDUOD SNYJUO/IYd IZE8T ‘suaydays snjzeubod SsnNyJUOliUd OP8T “‘UOSUDHF S17NeD SNYJUO/IYd L(ZO8T ‘JSsOyUaAPIN) SNLeUOGIED SNYJUO[IYd eUILJUO}IUd eUILUO|IUd eUILJUO}IUd eUILUOHIUd eUILUO}IUd eUILJUO}IUd eUILUO}IUd eUILUO|IUd eUILUO}IUd eUILUO}IUd eUILUO}IUd eUILUO}!Ud eUILUO}IUd eUILUO}IUd eUILJUO}IUd eUILUO}IUd eUILJUO}IUd eUILUO}IUd eUILUOIUd eUILUOIUd eUILJUO|IUd eUILUO|IUd eUILUO}IUd eUILUO|IUd eUILJUO}IUd eUILUOH!Ud eUILJUO}IUd eUILUO}IUd eUILUO}IUd eUILUO}IUd eUILJUO}IUd lujul|Aydeys jujul|Aydeys julul|AYydeys lujuljAydeys lujul|AYdeys lujuljAydeys julul|Aydeys jujul|Aydeys lujul|AYdeys lujul|Aydeys lulul|Aydeys jujul|Aydeys lujul|AYdeys julul|Aydeys lujul|Aydeys lujul|Aydeys julul|Aydeys juluI|AYydeys lujul|Aydeys jujul|AYdeys julul|AYydeys juIuI|AYydeys lujul|AYdeys lujul|Aydeys juluI|Aydeys lujul|Aydeys lujuljAydeys julul|Aydeys lujul|AYdeys juju|AYdeys lujul|Aydeys aeutuljAydejys aeurulAydeys aeuruljAydeys aeutulAydeys aeutuljAydeys aeuruljAydeys aeuruljAydeys aeurulAydejys aeutuljAydejys aeuruljAydeys aeuruljAydeys aeutulAydeys aeutuljAydeys aeuruljAyudejys aeuruljAydejys aeurulAydeys aeuruljAydeys aeuruljAydeys aeuruljAydeys aeuruljAydejys aeuruljAydejys aeuruljAydeys aeutuljAydejys aeutuljAydeys aeuruljAydeys aeutulAydeys aeutuljAydeys aeuruljAydeys aeuruljAydeys aeutulAydeys aeuruljAydeys NO. 53, 2019 BANISTERIA 66 WNSN Ul NAV-VA WNSN ul NAV-VA JIdA Ul NAV-VA WNSN ul NAV-VA HNWA ul NAV-VA dWM9 ul NAV-VA HNWA Ul NAV-VA AAV ul NAV-VA dWMD U! Hdus-VA iVA iVA VA iVA iVA iVA GW GW dW aw Od od Od od od TT0Z ‘UOIMAN sNssiUajae/d sNnIeIPAJE/d (O€ST ‘WiayiouUep|) sNbuojee/d snzeIpAIe|d (ST6T ‘AeseD) wnuould snze1pAjeld (OP8T ‘UOSYIIA) SNIBSAW SNIeIPAIeld (ZO8T ‘JSJOYUSARID) sNSO/NIeW SNIeIPAJC|d (O€8T ‘WIdYOUUeW)) SPE/NIELWLL SNICIPAJEld (ZO8T ‘JSJOYUSAPID) sOJeSSOY SNICIPAJE/d (LOST ‘Sniduge4) smelolway snze1pA}e/d (6€8T ‘UOsuUy) SUe;NxA SNIeIPAIE/d (€9gT ‘ayU0De7 *1°[) SaLwO2 sNIeIpAeld (ZO8T “JSJOYUSALID) SIAZGOLIEULID SNILIPAE/d (6€8T ‘UOSUdy) snsou/byed snzespAIe/d (ZO8T “JSIOYUSACID) SNVE/NBUID SAISa/OYJUO (€98T ‘9JU0D97 “1°f) sedipe (Snulagesed) snIeUayJOUIG x(ZO8T ‘JSIOYUSACIN) SNSO/IA SNSO/IXELU SN/IYAOILD (ST6T ‘Aased) SnaIUIBIA (Snyeyousiq) smipenQ bPbST ‘JOWIDUS|a,| "J*4 SNeUILA} (SruNesosIIp)) snIpan? TZ8T ‘WOH snaejads snaejads (sninesosolp)) snipen?) (908T ‘JSdOYUdAeID) sNULBaJad (sninesolp) snipanOE A(ZO8T ‘weysue;|) snujawosa (sninesowolp)) snipan? (ZO8T ‘JssoyUaAesD) syjorHey (sripand) snipanQ x(€€8T ‘suaydays) syjoany (snijydey) snipanQ H(Z6ZT ‘snidiiqe4) snpibjry (sninesoiolp) srupanQ ZS8T ‘Wiayiauuep| sazseboiypia (snunesosoip) snipeanQ? (908T ‘IsJOyUaAeID) snujondes (snyeYyoysiq) snipenO x(€P8T ‘wisysouuep|) seibeyd snyonuoipanQ I(ZO8T ‘JSIOYUSACID) SWEQUAA SNYJUO/IUYd L(68ZT ‘\IN4Aed) SUBLIEA SNYIUO/IYd eulul|Aydeys eulul|Aydeys eulul|Aydeys eulul|Aydeys eulul|Aydeys eululjAydeys eulul|Aydeys eululjAydeys eulul|Aydeys eulul|Aydeys eululjAydeys eulul|Aydeys eulul|Aydeys eulul|Aydeys eulul|Aydeys eullpand eullpand eUllpand eullpand eullpand eullpand eullpand eullpand eullpand eullpand eullpand PUILUO|IUd PUILUO|IUd julul|Aydeys julul|Aydeys julul|AYydeys julul|AYydeys lulu Aydeys juluI|AYydeys juluI|AYydeys lujul|Aydeys lulul|Aydeys julul|AYydeys julul|Aydeys julul|Aydeys lujul|Aydeys julul|Aydeys julul|Aydeys lujul|Aydeys lujul|AYdeys julul|Aydeys julul|Aydeys lujul|AYdeys lujul|AYdeys julul|AYydeys jujul|Aydeys lujul|Aydeys lujul|AYdeys julul|Aydeys juIuI|AYydeys lujul|AYdeys aeuruljAydejs aeuruljAydej}s aeuruljAydej}s aeuruljAydeys aeululjAydeys aeuruljAydej}s aeuruljAydej}s aeuruljAyde}s aeuruljAydej}s aeuruljAydej}s aeuruljAydej}s aeuruljAyde}s aeuruljAydej}s aeuruljAydej}s aeuruljAydej}s aeuruljAydej}s aeuruljAydej}s aeuruljAydej}s aeuruljAyde}s aeuruljAydeys aeuruljAydej}s aeuruljAydej}s aeurulAydeys aeurulAydeys aeuruljAydej}s aeuruljAydej}s aeuruljAydej}s aeuruljAydej}s 67 BRATTAIN ET AL.: ROVE BEETLES HM) ul €{V-VA dWM9-VA dWM9-VA OIdA ul NAV-VA dWM9S ul NAV-VA dWM9 ul NAV-VA WNSN Ul NAV-VA (pest ‘Aases) syiqewe snua7zs pggtT ‘Aase) /aIE/2 SNUAIS HG6LT ‘JAIN|O “D"V) SHeaul/ snujoyjueX (O88T ‘8}U0D27 “1'C) SluUadugns snuapojsiua]s LLZ8T ‘Jeyua||A5) snzezunaiurnued snyjeydoreyd L(8S8T ‘Aysinyos}oW|) s/uUadipyjed snjjeydoreyd Z86T ‘CURLS Mayayy sazde/qrxO (PZ6T ‘UPLUION) /SIAeD Sazaa/GAxO Q06T ‘AeseD s/uUadIpLIN] snigopny/ (OEst ‘Aes) sn/eyde? snigopny (ZS8T ‘asyoes) sryjisnd snudX{yoay (6€8T ‘UOSydZ) srunosqgo snudAyoay/ (906T ‘Aese>) sdouejaw snudAyoay (€88T ‘JaIAIANG) saquooay snudAyoayy (oggt ‘Aes) snjewey snudkyoay (O88T ‘28U0D97 “1'[) sdeaiasry snudXyoayy (906T ‘Aased) sy/bey snudkyoay (ZO8T ‘JSJOYUSAeID) snsauiua snudAyooy (6E8T ‘UOsYoy) smyenuaye snudkyooy/ (€98T ‘aqU0De7 “1'C) sY/oaIbuO/ SapOseYDOYIT L(EEQT ‘suaydays) snysnd snupejde7 L(OZZT “AINW "4°O) SloonIeY snudKyoIAD (Z8ZT ‘sniouqe4) snpibjny snuajdoineg 1(906T ‘saneyutag) Vayyuim (epayoerey) snibse | L(ZO8T ‘JsHOyUaAeID) vaje (sn/bse,) snibse (ZO8T ‘JSsOYUSAI) SMeUOZ SNILIPAIC|d (6Z8T “WIOH) SNUEDLIA snIeIpPAIe/d (ZO8T “JSIOYUSACID) SNAIC/OIA SNILIPAJE/d (ZO8T ‘JSIOYUSALID) SNSOJUALUIO] SNICIPATC|d eulul|Aydeys eululjAydeys eulul|Aydeys eululjAydeys eulul|Aydeys eulul|Aydeys jJuJUNjoU,UeX JuJUNjoU,UeX JuJUNjoU,UeX JuJUNjoU,UeX JUJUNjoU,UeX JuJUNoU,UeX jJujUsjou,UeX juJUJoU,UeX jJuJUNjoUJUeX JujUNjouyueX juUJoUuJUeX juJUJoUu,UeX JuJUNJoUu,UeX JuIUNJoU,UeX jJuJUNjoU,UeX JuJUNjoUu,UeX jJujUsjoujUeX JuJUNJoUu,UeX juJUNjoUu,UeX JuJUNjoUyUeX juJUNjoUyUeX julul|Aydeys juluI|AYydeys juluI|Aydeys julul|Aydeys julul|Aydeys juluI|Aydeys SPUIUDIS SEUIUDIS aeuruljAydeys aeuruljAydeys aeuruljAyudeys aeuruljAydejys aeuruljAyudeys aeutuljAydeys aeutuljAydeys aeuruljAydeys aeurulAydeys aeutuljAydeys aeutuljAydeys aeurulAydeys aeutuljAydeys aeutuljAydejys aeuruljAydeys aeutulAydejys aeutuljAyudeys aeuruljAydeys aeuruljAydejys aeutuljAydeys aeurulAudeys aeuruljAydejys aeutuljAydeys aeutuljAydeys aeuruljAydeys aeutuljAydejys aeutuljAydeys NO. 53, 2019 BANISTERIA 68 dWM9-VA aw GW GW GW dW aw GW Od od od TZ6T ‘ZYINd 4ebyas SsNuAIS EQET ‘a}U0DET “T' VO/oDIWWAS SNUAIS (p8gt ‘Aased) SNZ/PUOIAL SNPPUOIAL SNUAIS OP8T ‘UOSsUHy smyep—uNd snuajs b8st ‘Aasey snzipnd snuays b88T ‘AdSeD SILOCO/ILU SNUAIS (pEgt ‘AdSe?) SEUOIPLIALU SNUAIS OZ6T ‘UeWION /Z77/ sNUaIS (pest ‘AeseD) suabny snuajzs SZ6T ‘YOluag “1 SN/NJELUI/ SNUAIS 886T ‘ZYINd VOD) SNUAIS OP8T “UOSUDHF SILUODIAeY SNUAIS TEST ‘AWS SeIOWWAY SNUAIS (pest ‘Aase>) sdouna snuajzs OP8T ‘UuOosYyoy snuaba snuajs p8gT ‘Aasen/edsip sNuUaIS T00Z ‘ZUINd SNINIASEp snua}s P8RT ‘AdSED SISUBIEMB/AP SNUBS OT6T ‘Janeyusag sisuaqn? snuajs (pest ‘Aase)) SNJe890I9 SNUAIS ZLE6L ‘ZUINd SNSSAIAWOD SNUAIS x€98T ‘8JU0DE] “T'C BWLWOD snualsS OP8T ‘UOSUIA SNUOJOD Ssnuals TEST ‘Aes vojod snuajs PERT ‘AdSeD BEUI/OJED SNUAS OST ‘UOSUDH sNsoyer snsoyer snuajys TZ6T ‘ZYINd SUOYISO/ED sNUAIS 6I6T ‘UPWION SN/ODILALD SNUAIS p8gT ‘Aasen snjzebyUalig snuals OP8T ‘UOSUDHA srynze snuays ObST ‘UOSYDZ suejnuue snuazs SEUIUDIS eUIUD}S SEUIUDIS SEUIUDIS SEUIUDIS SeUIUD}S SEUIUDIS SeUIUDIS SEUIUDIS EUIUDIS SEUIUD}S SeEUIUDIS SEUIUDIS SseUulUaIS SEUIUDIS SeUIUDIS SEUIUD}S SEUIUD}S SEUIUDIS EUIUDIS SEUIUDIS SEUIUDIS SeUIUD}S SEUIUDIS SeUIUDIS SeUIUDIS SEUIUDIS SEUIUDIS SEUIUD}S EUIUDIS SeUIUDIS 69 BRATTAIN ET AL.: ROVE BEETLES HM) Uu! HM9-VA dWM9-VA dWM9-VA dWM9-VA iVA iVA GW GW Od Od OT6T ‘Heqnuds *y 1g Janeyusag /WOY snuodojaahp T66T ‘|leqdwe, sisuapuoy snuodojaahp/ EQ8T ‘A}U0DE] “]'[ S/OSUOD SNIOdOJBIAp/ 6E8T ‘UOSYOA SNUeIELUe SNIOdOJAIA/ (PhST ‘JOWIAYS|eW ‘4"4) SIISNUAA SNIIDCJOYI UOYJIPJ1OT (ZZ8T ‘UJOH) sozsaenb UOYzIpPI07 (ZERT ‘AeS) sNjajOsqo UOLJIPsOT Z86T ‘Ileqduiey sy/gejou UOLYPIOT (ZO8T ‘JSJOUUSACIN) ABIL UOYJPJOT (pr6T ‘UDIEW) Aeyey YoYyppso7 (Gggt ‘Aase>) ses UOY/JIPIOT (ZO8T “JSIOYUSALID) SJILID UOLYJIPJIOT 0002 ‘AxINYDS Wages UOY}P1O7 (908T ‘JSIOYUSARID) SLIEYIXe UOYJPIOT Z86T ‘||eqduieD snuenjpejedde UOYP/OT (ZZ8T ‘UWIOH) sn2gUe UOYJPIOT (ZT6T ‘Janeyusag) LWADLIBIA PLUOSOUYIST T66T ‘lleqdwe Wayns ELIOSOUYIST x(908T ‘JSsOYUdAeI) LUNpPIPUAa/As ELUOSOUYIST T66T ‘||eqdwe> /a7U0Ia/ ELIOSOUYIST (€98T ‘a}U0De7 “T'C) a/oDINey ELIOSOUYIST TOOZ ‘UeWHAH sreuaLya siIeydiez (6€8T ‘UOsYoIZ) smepiUp sioeYydie? COQT ‘A]U0De7 “1° sNaIe7Sa) sNIodoAg COST ‘BU0De] “T'[ suaIsajyn sruodoAig E66T ‘|eqdwes seuvejaws syeydosig LOEST “WilayiouUeW s7e;nbuld snigoqyyjog bEQT ‘AdSED BEILIGIIA SNUAIS b8gT ‘AdSED SNLIDIA sNUAS TEgt ‘Aes snaibAjs snuajs lULIOdO}@dA|} 1ULIOd0}adA|| lULIOd0}dA|| I1ULI0d0}@dA|} 1ULIOdO}dAW 1ULIOd0}a@dA|J lULIOd0}adA|J 1ULI0d0}adA|} 1ULIOd0}dA|} 1ULIOdO,SdDA|) lULIOd0}dA| I1ULIOdO,@dA|} lULIOd0}SdA| IULIOd0}adA|\J 1ULIOdO}@dA|} lULIOdOJSdA} lULIOdO}SdA|W lULIOd0}adA|J 1ULIOdO,dA|J l1ULI0d0}adA|J lULIOdO}SdA|} 1ULIOdO}SdA|} lULIOd0}dAW 1ULIOdO,@dA|} I1ULIOd0,dA|) lULIOd0}adA|) lULIOd0 dA) aeutodAyoeL aseutodAyoeL aeutiodAyoeL aseunodAyoeL aeutodAyoeL aseutodAyoeL aeutiodAyoeL aseunodAyoeL aseutiodAyoeL aeutiodAyoeL aseutiodAyoeL aseutiodAyoeL aseutiodAyoeL aeutodAyoeL aseutodAyoeL aseutodAyoeL aseutiodAyoeL aseutodAyoeL aeutiodAyoeL aseutiodAyoeL aseutodAyoeL aeutiodAyoeL aseutodAyoeL aeutiodAyoeL aseutodAyoeL aeutiodAyoeL aeutodAyoeL SeUIUD}S SeUIUD}S SPUIUDIS NO. 53, 2019 BANISTERIA 70 dWM9-VA GW od Od od ZO8T “JSIOYUSACID SNIELIQUIY SNUIYIE 1 LL8T ‘UJOH SISUQPEURD SNUIYIE 1 *OE8T ‘UOSUDIF SyeSeg SNUIYIE | 6E8T ‘UOSUDHF SUeyIXe SNUIYIC | LL81 ‘UJOH SnpUappe snulyoe J (GggT ‘AaseD) 40/02ISIAA snjlydopadas (S6gt ‘Aase?) sadiza0jan snjiydopadas I(Z6ZT ‘sniouge4) snazej7saz snjilydopadas (ZZ8T ‘UWIOH) snduOS snrjitydopadas (ZZ8T ‘UJOH) snynvued snjiydopades (zEegt ‘Aes) snzido snjitjdopedas (gst ‘Aese)) $19/NI90 snjil{dopadas (G6gt ‘Aase)) va2eW Snjiy~dopadas 1(8SZT ‘Snaeuurq) snavojy snyilydopades O/6T ‘W!eqdue Maazy snjily¥dopadas O/6T ‘I|eqdue 7 Wsoy snjiydopadas (6st ‘Aesed) syiqap snjiydopades 9/6T ‘lleqduiey syepiuaja snjilydopades (ZO8T ‘JssoyUeAesD) snsse/D snjiydopedes (6E8T. ‘UosYoLA) sryNjoUID snyydopades T00Z ‘UBWHAH Wjegdues snjilydopadas 9/6T ‘I|eqdueD snajdhyoeig sryiydopades (6E8T ‘UOsUdI4y) syeseq snjiydopades (898T ‘PlOJeH 19 JABUILWLUAS) /OZENOS SNUIYICEJOPHIN (€Z6T ‘Ieqdwed) soy snuILIe}0pHIN (zest ‘Aes) sninatyuan sniodosdoj CQO8T ‘8]U0D27] “T'[ SMNae/ srUOdo/doOD H(Z9ZT ‘snaeuur)) saployays eaflD CQQT ‘AaqU0DeET “TC SN/NPION sruodojaahp luuOdAype | luuOdAype | luuOdAYpe | luuOdAype | luuOdAyoe | luOdAYpe | luLOdAyoe | luLOdAyYoe | luuOdAype | luuOdAype | luuOdAype | luuOdAyYpe | luuOdAype | luLOdAyoe | luLOdAYpe | luuOdAyoe | luuOdAYpe | luOdAYpe | luuOdAyYpe | luuOdAype | luLOdAype | luLOdAype | luuOdAype | luLOdAYpe | luLOdAYoe | luuOdAyoe | luuOdAYpe | luuOdAype | lULIOd0}adA|} aseutiodAyoeL aseutiodAyoeL aseutodAyoeL aeutodAyoeL aeutodAyoeL aeutiodAyoeL aseunodAyoeL aseutiodAyoeL aseutiodAyoeL aeutiodAyoeL aseutiodAyoeL aseutiodAyoeL aeutiodAyoeL aseutiodAyoeL aeutodAyoeL aeutodAyoeL aseutiodAyoeL aseutiodAyoeL aseutodAyoeL aseutiodAyoeL aseutiodAyoeL aeutodAyoeL aseutodAyoeL aeutodAyoeL aseutiodAyoeL aseutodAyoeL aseutiodAyoeL aseutiodAyoeL aseutnodAyoeL 71 BRATTAIN ET AL.: ROVE BEETLES dWM9-VA LOTST ‘Jeyual|Ad) sywoayid eAYdOYOL) L(T8ZT ‘sniouqe4) sninpaiu (sniodjed) sniodAyoe | 6E8T ‘UOsSYyoHA snueU (snuodAyoe 1) sniodAyre J 6/6T ‘l|aqdwe sazuoza/ (sruodAyoe 1) sruodAyoe | *ZERT ‘Aes snsozol (sniodAyoe 1) snuodAyre J Z/8T ‘UJIOH suebaa (snuodAyoe 1) sniodAyoe J 6E8T ‘UOSUudHy sadioid snuiy7e | €/6T ‘|]eqdwes snusuiLU SNUuIYIe J ZO8T ‘JSIOYUSALID SN/VOULUALU SNUIYIC | OP8T ‘UOSUDHA SPUN] snulyze | PHST ‘JOWIOYS|aW “J'4 SMJEGLUI/ SNUIYOE | (zEgt ‘AesS) suuadnUny SNuIYyre | luLOdAYoe | luuOdAYpe | luuOdAype | luOdAYe | luuOdAype | luuOdAYpe | luOdAype | luLOdAYoe | luLOdAYpe | luLOdAYoe | luLOdAYoe | aeurAydousu aeutnodAyoeL aseutiodAyoeL aeutiodAyoeL aeutiodAyoeL aeutiodAyoeL aseutniodAyoeL aeutiodAyoeL aeutodAyoeL aeutiodAyoeL aeutodAyoeL aseutodAyoeL Banisteria, Number 53, pages 72—77 © 2019 Virginia Natural History Society Probable Cerulean Warbler x Northern Parula Hybrid in Rockbridge County, Virginia in April 2019 Richard A. Rowe Department of Biology Virginia Military Institute Lexington, Virginia 24450 RoweRA@vmi.edu Lucinda M. Rowe Roanoke, VA ABSTRACT We report our sighting of a probable Cerulean Warbler (Setophaga cerulea) x Northern Parula (Setophaga americana) hybrid that was located in Rockbridge County, Virginia on 20 April and 28 April 2019. Superficially, the bird resembled an after second-year male Cerulean Warbler, but it had several plumage characteristics of an after second-year Northern Parula. Additionally, the hybrid sang a Northern Parula song. This would be the first record of this hybrid in Virginia. Keywords: Cerulean Warbler, hybrid, Northern Parula, wood warbler. INTRODUCTION The wood-warblers (family Parulidae) are a well- known group of small insect-eating birds. Lovette et al. (2010) have presented a recommendation for revising the relationships among the parulids. They used molecular techniques to assess phylogenetic relationships and, of interest to our paper, they show that Cerulean Warblers and Northern Parulas are very closely related. Recently, Trimbath et al. (2019) reported on a genetic analysis of Cerulean Warbler x Northern Parula hybrids from northeastern Ohio. They showed that the two individuals had the same Cerulean Warbler dam and a Northern Parula sire. The presence of hybrids poses interesting questions with respect to speciation and evolution. Hybridization can be due to a number of reasons. Most commonly, there exists a hybrid zone where the ranges of two closely related species who only recently diverged and have retained many similarities interbred. In birds, hybridization has been reported in many families (Cockrum, 1952) and interestingly there is a high rate of hybridization in waterfowl (Ottenburghs et al., 2016). The presence of hybrids in the Parulidae has long been recognized; hybrids produced between Golden-winged Warblers and Blue-winged Warblers, Brewster’s Warbler or Lawrence’s Warbler being the most recognized (Short, 1963). A number of other parulid hybrids have been reported, for example: Northern Waterthrush x Blackpoll Warbler (Short & Robbins, 1967); Black-and-white Warbler x Cerulean Warbler (Parkes, 1978); Orange-crowned Warbler x Nashville Warbler (Ralston et al., 2015); Chestnut-sided Warbler x Magnolia Warbler (Burrell et al., 2016); Cerulean Warbler x Black-throated Blue Warbler (Delancey et al., 2019); Cerulean Warbler x Northern Parula (Trimbath et al., 2019). In this paper, we report on a Cerulean Warbler x Northern Parula hybrid that was discovered in Rockbridge County, Virginia in April 2019. METHODS Location Rockbridge County is located in southwestern Virginia and is bordered by the Blue Ridge Mountains on the east and Appalachian Mountains on the west. ROWE & ROWE: HYBRID WARBLER 73 Geographically, Rockbridge County is in the Great Valley of Virginia, and it is situated at the southern end of the Shenandoah Valley. The Maury River, the principal waterway within the county, flows into the James River near the southeastern boundary of the county. The location for the hybrid was along the Maury River south of the city of Buena Vista and along River Road, which parallels the Maury River for several miles. The hybrid was found in an area (37°40'28.33"N; 79°25'31.86"W) with Sycamore (Platanus occidentalis) trees bordering a small creek and wet area within 100 meters of the Maury River. Equipment The hybrid was photographed using a DSLR camera with a telephoto lens. Sound recordings were made using a Telinga Pro parabolic dish, Sennhieser microphone (MKH 20) and a Fostex FR-2 digital field recorder. In addition, sound recordings of a Cerulean Warbler (ML 85132) and Northern Parula (ML 53302) were obtained with permission from the Macaulay Library at the Cornell Lab of Ornithology. Spectrograms from the hybrid, Cerulean Warbler, and Northern Parula songs were generated using Raven Pro 1.4 (Bioacoustics Research Program, The Cornell Laboratory of Ornithology). RESULTS AND DISCUSSION On the morning of 20 April 2019, while birding along River Road and the Maury River in southern Rockbridge Co., Virginia, we discovered the hybrid. The bird was first located by its song, which was recognized by one of us (RAR) as a Northern Parula. As a result, we began looking for a Northern Parula. The bird continued to sing and we were able to locate it in trees. Once we saw the bird, we changed our identification to a Cerulean Warbler based on its plumage presentation. We were somewhat confused, however, because we were hearing a Northern Parula but seeing a Cerulean Warbler. In an effort to get better looks at the bird, we played vocalizations for both Cerulean Warbler and Northern Parula from iBird (version 10.06). At the time, we felt that the hybrid tended to respond to the Cerulean Warbler vocalizations better than the Northern Parula. We photographed the bird (Fig. 1) and later when we reviewed our photographs we realized that there were some inconsistencies with the plumage and our Cerulean Warbler identification. As a result, we returned to the location on 28 April and quickly relocated the bird. On that date, we had the assistance of another observer and were able to record the hybrid singing while confirming that 1t was the hybrid singing. We used Raven Pro to create spectrograms of the hybrid’s vocalizations (Fig. 2). Additionally, we experimented with playing vocalizations of both the Cerulean Warbler and Northern Parula from iBird and noting the responses of the hybrid. It was clear that the hybrid gave a greater response to the Cerulean Warbler call by flying from perch to perch on different trees singing. Description of hybrid Overall, the hybrid (Fig. 1) resembled a Cerulean Warbler exhibiting a blue dorsum and head with a blue breast stripe, white belly and throat, weak blue spotting along the flanks, two prominent white wingbars, and white undertail coverts. The hybrid lacked the yellow coloration seen on the throat and breast of a Northern Parula as well as the orange wash seen below the breast band. The upper mandible was black and the lower mandible was gray, which is consistent with a Cerulean Warbler (Northern Parulas have a black upper mandible and a yellow lower mandible). The hybrid had a clear split eye-ring (a Northern Parula characteristic but can be seen in immature Cerulean Warblers), and a completely blue-gray head and neck instead of the sky- blue coloration seen in Cerulean Warblers. The hybrid lacked the white neck band that extends between the nape and cheek (seen in Cerulean Warblers at all ages), which is consistent with a Northern Parula. Also, some of our photographs appear to show a slight green tinge to the dorsum between the wings (Northern Parulas have a distinct greenish triangle between their wings), but no black longitudinal stripes were visible (a Cerulean Warbler characteristic). In addition, the photographs do not show the distinct black, longitudinal stripes seen in adult Cerulean Warblers. Unfortunately, none of the photographs provide a clear view of the back of the bird. We recorded a total of seven minutes of the hybrid singing. A review of the spectrograms from our recordings were all consistent with the vocalizations of a Northern Parula. A representative spectrogram from the vocalizations recorded from the hybrid is presented in Fig. 2. The hybrid’s spectrogram is consistent with the description of the type A Northern Parula song (see Fig. 3) described by Moldenhaur (2012) as containing a preliminary trill ending with a buzzy trill and inverted chevron terminal note. Cerulean Warblers (see Fig. 4) have an introductory set of longer notes followed by a middle section of shorter notes and ending with a final buzz (Buehler et al., 2013) but they lack the inverted chevron of a Northern Parula. We compared the vocalizations that we recorded to spectrograms from Northern Parula (Fig. 3, ML 53302) and Cerulean Warbler (Fig. 4, ML 85132) and found that the vocalizations from the hybrid are consistent with the 74 BANISTERIA NO. 53, 2019 Fig. 1. Photographs of the Cerulean Warbler x Northern Parula hybrid taken on 20 April 2019 in Rockbridge Co., Virginia (photographs by R. A. Rowe). ROWE & ROWE: HYBRID WARBLER 75 Le \ . , y 2 |e a f.: Wa Yala. shit us my ih +. >| : ny) ‘ /) if TATE OA. ; j 0 ae : ipa \ , AT \ JAR Ab it 9 f SO i Di i) Me (na Hie i a | 2] vei ’ Wi i" | } j hy bd thet War vi i NOM y i 1," ‘tae et) Wy ty | ly . NY, wat A) Ver hilt a Ufa det Lule " Hi yy if Min Wl ’ i) NW 4 Hae 4 Me) yal 4, n Wii , 0.000 kHz f 1 , , ify x 2.4 30 0.2 0.4 0.6 0.8 1 1.2 1.4 1.6 1.8 2 2.2 2.6 2.8 3 Fig 2. Spectrogram from Cerulean Warbler x Northern Parula hybrid recorded on 28 April 2019 along the Maury River south of Buena Vista, VA. Spectrogram created using Raven Pro 1.4 (Bioacoustics Research Program, Cornell University). Fig. 3. Spectrogram from a Northern Parula (ML 53302, with permission Macaulay Library at Cornell Lab for Ornithology) created using Raven Pro 1.4 (Bioacoustics Research Program, Cornell University). 12 (yl i A) Yi! i iN ae t t)) | ; vy), aa mL) n| 1 2 1 Vee Sake JUL A 0.000 bi Mtlad Wet EEL Ltn ee al a a ee ee Widiu whe aed Ne eee | Metz $0 0.2 0.4 0.6 0.8 1 a fe 1.4 1.6 1.8 2 2.2 2.4 Fig. 4. Spectrogram from a Cerulean Warbler (ML 85132, with permission Macaulay Library at Cornell Lab for Ornithology) created using Raven Pro 1.4 (Bioacoustics Research Program, Cornell University). 76 BANISTERIA vocalizations of a Northern Parula (Fig. 3) and not a Cerulean Warbler (Fig. 4). In addition, we compared our recordings of the hybrid bird to those reported by Trimbath et al. (2019) and note that there is a striking similarity between the two hybrid’s vocalizations. If we apply the interpretations of the vocalizations reported by Trimbath et al. (2019) to our recordings, we suspect that our hybrid had a Cerulean Warbler dam and a Northern Parula sire. There are only a few reports of Cerulean Warbler x Northern Parula hybrids. Pooth & Johnson (2004) reported a suspected hybrid in Dutchess Co., New York, and Nirschl (2004) reported a hybrid in Toledo, Ohio. Lindsey & Vezo (1995) described a_ suspected interbreeding between a female Cerulean Warbler and a male Northern Parula on Long Island, New York. Trimbath et al. (2019) documented genetic hybridization in two individuals captured in northeastern Ohio. Reports of suspected hybrids have been submitted as photographs and/or sound recordings via eBird to the Macaulay Library at the Cornell Lab of Ornithology. These reports are from various locations with multiple reports from some locations submitted via eBird. In total, there appear to be five individuals being described (Kane, Illinois with photographs or recordings from April and May 2019 at the same location and most likely the same bird: see for example - ML 156727611, 6 May 2019; ML 155845381, 4 May 2019; ML 155532821, 1 May 2019; ML 63231511, 14 May 2017; Cape May, New Jersey ML 139279761, 4 May 2009; ML 63156701, 13 May 2009; Muscatine Iowa ML 59905421, 13 May 2017, Marion Co (Indianapolis), Indiana ML 27567461, 21 April 2016). It is interesting to note that photographs of proposed Cerulean Warbler x Northern Parula hybrids available at the Macauley Lab show plumage characteristics that are remarkably similar to the bird that we located. In addition to the Cerulean Warbler x Northern Parula hybrids, other hybrids involving Cerulean Warblers have been reported: Cerulean Warbler x Black- and-white Warbler (Parkes, 1978) and Cerulean Warbler x Black-throated Blue Warbler (Delancey et al., 2019). Hybrids involving Northern Parulas include: Northern Parula x Yellow-throated Warbler, known as Sutton’s Warbler (Anich et al., 2012) and Northern Parula x Yellow Warbler (Graves, 1993). Hybridization in the family Parulidae is not uncommon as shown by the many reports of hybrids in this group. The bird we located exhibited plumage characteristics that are consistent with both Cerulean Warblers and Northern Parulas. Additionally, while the bird superficially resembled a Cerulean Warbler, it sang a Northern Parula song. A few reports of a Cerulean Warbler x Northern Parula hybrid can be found in the NO. 53, 2019 literature or at the Macaulay Library. In these cases, the birds show a mix of plumage characteristics and primarily resemble a Cerulean Warbler. When the song was described or recorded, the hybrids tended to sing a Northern Parula song. Without genetic analysis, we cannot confirm that the bird we discovered was truly a hybrid, but we believe the evidence is compelling that this bird is a Cerulean Warbler x Northern Parula hybrid. ACKNOWLEDGEMENTS We would like to thank Ruth Rowe for her assistance in recording the vocalizations of the hybrid and the Macaulay Library at the Cornell Lab of Ornithology for permission to use sound files. Funding that supported this research was received from the Virginia Military Institute’s Grants-in-Aid of Research and the Biology Department. LITERATURE CITED Anich, N. M., J. A. Swelstad, & B. Pierce. 2012. The first report of Sutton’s Warbler (Northern Parula x Yellow- throated Warbler) in Wisconsin. Passenger Pigeon 74: 149-155. Burrell, K. G. D., J. H. Skevington, S. Kelso, M. V. A. Burrell, D. L. Leclair, & S. A. Mackenzie. 2016. Previously undocumented hybrid new world warbler (Setophaga pensylvanica x S. magnolia) captured at Long Point, Ontario. Wilson Journal of Ornithology 128: 624-628. Buehler, D. A., P. B. Hamel, & T. Boves. 2013. Cerulean Warbler (Setophaga cerulea), version 2.0. In The Birds of North America (A. F. Poole, Editor). Cornell Lab of Ornithology, Ithaca, NY, USA. https://doi.org/10.2173/bna.511 (accessed 10 July 2019) Cockrum E. L. 1952. A check-list and bibliography of hybrid birds in North American north of Mexico. Wilson Bulletin 64: 140-159. Delancey, C. D., G. J. MacDonald, & K. Islam. 2019. First confirmed hybrid pairing between a Cerulean Warbler (Setophaga cerulea) and a Black-throated Blue Warbler (Setophaga caerulescens). Wilson Journal of Ornithology 131: 161-165. Graves, G. R. 1993. A new intergeneric wood warbler hybrid (Parula americana x Dendroica coronata) (Aves: Fringillidae). Proceeding of the Biological Society of Washington 106: 402-409. ROWE & ROWE: HYBRID WARBLER 77 Lindsay, P., & T. Vezo. 1995. Cerulean Warbler and Northern Parula tending the same nest. Kingbird 45: 160-161. Lovette, I. J., J. L. Perez-Eman, J. P. Sullivan, R. C. Banks, I. Fiorentino, S. Cordoba-Cordoba, M. Echeverry-Galvis, F. K. Barker, K. J. Burns, J. Klicka, S. M. Lanyon, & E. Bermingham. 2010. A comprehensive multilocus phylogeny for the wood- warblers and a revised classification of the Parulidae (Aves). Molecular Phylogenetics and Evolution 57: 753— 770. Moldenhauer, R. R., & D. J. Regelski. 2012. Northern Parula (Setophaga americana), version 2.0. In The Birds of North America (A. F. Poole, Editor). Cornell Lab of Ornithology, Ithaca, NY. https://doi.org/10.2173/bna.215 (accessed 10 July 2019) Nirschl, R. (2004). A possible Northern Parula x Cerulean Warbler hybrid in Toledo, Ohio. Ohio Cardinal 27: 164-168. Parkes, K. C. 1978. Still another parulid intergeneric hybrid (Mniotilta x Dendroica) and its taxonomic and evolutionary implications. Auk 95: 682-690. Pooth, C., & R. Johnson. 2004. An unusually plumaged Cerulean Warbler singing a Northern Parula song in Dutchess County, NY. Kingbird 54: 314-316. Ottenburghs, J., P. van Hooft, S. E. van Wieren, R. C. Ydenberg, & H. H. T. Prins. 2016. Hybridization in geese: a review. Frontiers in Zoology 13: 20. DOI 10.1186/s12983-016-0153-1 Ralston, J., D. A. Ermacor, & J. J. Kirchman. 2015. A new wood warbler hybrid (Oreothlypis celata x Oreothylpis ruficapilla) from the Adirondack Mountains, New York. Wilson Journal of Ornithology 127: 21-28. Short, L. L., Jr. 1963. Hybridization in the wood warblers Vermivora pinus and JV. chrysoptera. Proceedings of the 13th International Ornithological Congress 147-160. Short, L. L., Jr., & C. S. Robbins. 1967. An intergeneric hybrid wood warbler (Seiurus x Dendroica). Auk 84: 534-543. Trimbath, R. J., A. W. Jones, & C. L. Brennan. 2019. Evidence for hybridization between Northern Parula (Setophaga americana) and Cerulean Warbler (S. cerulea). Wilson Journal of Ornithology 131: 615— 624. Banisteria, Number 53, pages 78-83 © 2019 Virginia Natural History Society First Records of the Neotropical Deer Ked Lipoptena mazamae Rondani (Diptera: Hippoboscidae) from Virginia Liberty Hightower, Nancy D. Moncrief, and Kaloyan Ivanov Virginia Museum of Natural History 21 Starling Avenue Martinsville, Virginia 24112 liberty.hightower@vmnh. virginia. gov ABSTRACT We report the first definitive records of Lipoptena mazamae Rondani, a hematophagous ectoparasite of cervids, from the Piedmont of Virginia based on recently collected specimens. The collection localities in Henry and Pittsylvania counties represent the northernmost known occurrence of this species in eastern North America. Keywords: eastern United States, ectoparasites, range extension, zoonotic diseases. INTRODUCTION Louse flies (Diptera: Hippoboscidae) are a small group of obligate hematophagous ectoparasites of birds and mammals represented in the world’s fauna by more than 210 species in three subfamilies (Ornithomyinae, Hippoboscinae, and Lipopteninae) (Dick, 2006). These unusual flies are easily recognized by their robust, dorsoventrally flattened bodies, “leathery” appearance, and highly modified antennae and mouthparts (Maa & Peterson, 1987). Most hippoboscids retain functional wings throughout their life, while members of Lipopteninae are either wingless or shed their wings upon locating a host. Lipopteninae are normally parasitic on cervids (Artiodactyla: Cervidae), while most members of the remaining two subfamilies primarily parasitize birds (Maa, 1969; Maa & Peterson, 1987). Both sexes readily take blood meals and some species are known, or suspected, vectors of etiological agents such as viruses, bacteria, protozoans, and helminths associated with a variety of conditions including bluetongue, West Nile fever, anaplasmosis, bartonellosis, —_ borreliosis, ehrlichiosis, pseudomalaria, and trypanosomiasis (Farajollahi et al., 2005; Small, 2005; de Bruin et al., 2015; Liu et al., 2016; Kelsey & Finch, 2018; Moreira et al., 2019; Skvarla & Machtinger, 2019). The pigeon fly, Pseudolynchia canariensis (Macquart), an obligate parasite of pigeons, is the definitive host and the only known vector of the protozoan Haemoproteus columbae Kruse (Moreira et al., 2019). The bluetongue virus is transmitted by the sheep ked, Melophagus ovinus (Linnaeus), to its main host the domestic sheep, Ovis aries Linnaeus (Small, 2005). In addition, RNA of the West Nile virus was recently detected in Jcosta americana (Leach) on birds of prey (Farajollahi et al., 2005). Although the actual transfer of pathogens by many of these flies is not completely understood, vertical transmission of Bartonella spp. recently has been shown between generations of the sheep ked, MZ ovinus, and the deer ked, Lipoptena cervi (Linnaeus), (Halos et al., 2004; de Bruin et al., 2015). The subfamily Lipopteninae includes three genera and 34 recognized species (Dick, 2006). Three native (Lipoptena depressa |Say], L. mazamae Rondani, and Neolipoptena ferrisi [Bequaert]) and two introduced (L. cervi and M. ovinus) species are known from the Nearctic (Maa, 1969; Dick, 2006; Skvarla & Machtinger, 2019). The Neotropical deer ked, ZL. mazamae, the introduced deer ked, L. cervi, and the sheep ked, M. ovinus, are the only members of Lipopteninae known to occur in eastern North America, the last two of which have been previously reported from Virginia based on verified records (Bequaert, 1957; Small, 2005; Skvarla & Machtinger, 2019). Lipoptena mazamae was first reported as occurring in Virginia by Kern (2003), HIGHTOWER ET AL.: NEOTROPICAL DEER KED 79 a record subsequently cited in Skvarla & Machtinger (2019), but neither study included examination of specimens from Virginia (M. J. Skvarla, pers. comm. to LJH, 4 February 2019; W. H. Kern, pers. comm. to LJH, 26 February 2019). Here we report the first definitive records of L. mazamae from Virginia based on recently collected specimens, and discuss the potential range overlap and interaction between L. mazamae and L. cervi. MATERIALS During routine processing of an adult female and a juvenile male White-tailed Deer, Odocoileus virginianus (Zimmermann), at the Virginia Museum of Natural History, the first author observed numerous dealate hippoboscid flies, which were collected, mounted, and subsequently identified as L. mazamae (Fig. 1). The female deer was collected on 1 January 2019, and the entire right rear leg (NDM 4488) was removed and placed in a plastic bag within 30 minutes of the animal’s death. The leg was brought to the Virginia Museum of Natural History and placed in a -20° C freezer on 2 January 2019. On 25 January, the leg was thawed and all of the skin was removed as part of procedures to prepare the bones as an articulated partial skeleton. During skinning, three female deer keds were found concealed in the white fur of the inner leg near the top of the calcaneus. It is unknown whether this was the total number of keds present on the host, as only the right rear leg was available for examination. The host from which these keds were extracted was collected on private property in Pittsylvania Co., Virginia approximately 14 km west of the City of Danville (36.60457°, -79.55312°). The male deer (NDM 4497) was collected on the morning of 9 October 2019, and the entire carcass was placed in a plastic bag within 12 hours of the animal’s death. The carcass was brought to the Virginia Museum of Natural History and placed in a -20° C freezer on the same day. On 19 November the carcass was thawed and the skin was removed as part of procedures to prepare the bones as a skeleton. During skinning, 82 deer keds (39 males, 43 females) were found, most of which were Fig. 1. Dorsal view of (left) dealate female Lipoptena mazamae (VMNH 110610) from Pittsylvania Co., Virginia; and (right) dealate male Lipoptena mazamae (VMNH 110619.1) from Henry Co., Virginia. Photographs by Lucy Treado, Virginia Museum of Natural History. 80 BANISTERIA concealed in the white fur of the inner rear legs and groin area. Due to poor preservation, only 75 keds (37 males and 38 females) were retained for deposition as specimens. The host from which these keds were extracted was collected on private property in Henry Co., Virginia approximately 8.5 km east of the City of Martinsville (36.67623°, -79.77738°). All specimens are deposited in the invertebrate collection at the Virginia Museum of Natural History (VMNH_ 110610-116012 from Pittsylvania Co., and VMNH 110619.1-110619.75 from Henry Co.). IDENTIFICATION Members of Lipopteninae are characterized by the fairly dense coverage of evenly spaced spiniform setae on the ventral surface of the thorax; the strongly concave posterior margin of the basal abdominal sternite (convex in N. ferrisi),; and the greater size of the basal abdominal sternite which is distinctly larger than the scutellum (Maa & Peterson, 1987). Among the three eastern members of Lipopteninae, M. ovinus 1s easily identified by its rudimentary wings, the absence of halteres, and the reduced eyes, which are each narrower than the antenna. The remaining two eastern taxa are separated based on the position of the median abdominal tergal plates (tergites 4—6 in males, 4—7 in females). In L. cervi these are evenly distributed over the abdomen, while they are crowded together in the posterior 1/3 of the abdomen in L. mazamae (Fig. 1; see also Figs. 1-5 in Skvarla & Machtinger, 2019). Keys to the Nearctic Lipopteninae genera and species are found in Maa (1965) and Skvarla & Machtinger (2019). DISCUSSION Lipoptena mazamae is an ectoparasite of White- tailed Deer, South American Red Brocket Deer, Mazama americana (Erxleben), Central American Red Brocket Deer, M. temama (Kerr), and Gray Brocket Deer, M. gouazoubira Fisher (Bequaert, 1942; Maa 1969). Incidental hosts include cattle, Bos taurus Linnaeus, Tayra, Eira barbara (Linnaeus), Peccary, Pecari tajacu (Linnaeus), Cougar, Puma concolor (Linnaeus), Pampas Deer, Ozotoceros bezoarticus (Linnaeus), and humans, Homo sapiens Linnaeus (Bequaert, 1942: Bequaert, 1957; Reeves et al., 2006; Graciolli et al., 2011; Shock, 2014). The parasite is commonly found on the legs, abdomen, and genital areas of the host (Samuel & Trainer, 1972). Deer populations may have infestation rates of 60-80%, with individuals sometimes harboring hundreds of keds (Bequaert, 1952; Samuel & Trainer, 1972; Kern, 2003). NO. 53, 2019 The known range of L. mazamae includes North, Central, and South America (Kennedy et al., 1987; Skvarla & Machtinger, 2019). South of the United States, this species occurs in Argentina, Bolivia, Brazil, Colombia, Costa Rica, Ecuador, French Guiana, Guatemala, Guyana, Honduras, Mexico, Nicaragua, Panama, Paraguay, Peru, Suriname, Trinidad, Uruguay, and Venezuela (Bequart, 1957; Maa, 1969; Carreno et al., 2001; Deem et al., 2004; Genoways & Timm, 2005; Romero-Castafion et al., 2008; Graciolli et al., 2011). It reaches as far south as Uruguay and Santiago del Estero in northern Argentina (Bequaert, 1942; Maa, 1969). In the United States, L. mazamae records based on specimens and photo vouchers have been reported from Alabama, Arkansas, Florida, Georgia, Louisiana, Mississippi, Missouri, North Carolina, Oklahoma, South Carolina, Tennessee, and Texas (Skvarla & Machtinger, 2019). The northernmost known localities of L. mazamae include St. Clair Co., MO in the West; and Anderson Co., TN and Iredell and Wake counties, NC in the East (Skvarla & Machtinger, 2019). Our records of L. mazamae reported herein may be an indication that this species is naturally expanding its range northward. Climate, and not the absence of its primary host the White-tailed Deer, appears to be the limiting factor for northward expansion of L. mazamae (Bequaert, 1954). Bequaert (1954) reported reduction of L. mazamae numbers, even to the point of complete absence, during harsh winters when frost was common and below-freezing temperatures were frequent. Given current long-term climate change projections, it is likely that L. mazamae will continue to expand its range northward (Collins et al., 2013). Alternatively, LZ. mazamae may have reached Virginia through human-mediated translocation of its host. White-tailed Deer were nearly extirpated from Virginia in the early 1900s (Handley & Patton, 1947). Between 1930 and 1950, at least 1,800 animals were translocated from Florida, Michigan, North Carolina, Pennsylvania, and Wisconsin into Virginia (Linzey, 1998). Deer from Florida, where the parasite has been present since at least the mid-1930s, could have harbored L. mazamae at the time of their relocation (Bequaert, 1935; 1942). Potential overlap of the geographic ranges of L. mazamae and L. cervi in Virginia and the surrounding areas was suggested by Skvarla & Machtinger (2019). Our records provide further evidence to support this possibility. Such a scenario represents an interesting ecological situation in which two species that have evolved on different continents might co-occur on the same individual host animal. We can only speculate regarding any potential interactions between the two HIGHTOWER ET AL.: NEOTROPICAL DEER KED 81 taxa, which may range anywhere from exploitation competition to avoidance. It is, however, likely that under such circumstances, the two taxa might exploit different areas of the host’s body, a situation previously observed in western deer keds and some ticks (Westrom & Anderson, 1992; Baer-Lehman et al., 2012; Skvarla & Machtinger, 2019). Actual geographical overlap between the ranges of the two species has not yet been documented. Lipoptena mazamae is currently only known from the Piedmont of Virginia while L. cervi has been reported from Craig, Montgomery, Shenandoah, and Wythe counties in the Blue Ridge and Ridge & Valley physiographic regions in the mountainous, western part of the state (Skvarla & Machtinger, 2019). Given the close proximity (<80 km) of their currently known ranges, it 1s possible that a direct physical overlap might occur via natural dispersal of the host and/or the parasites. The annual home ranges of adult male deer in Virginia have been estimated at 2.5 km/?, with dispersing young moving 6.4 km on average (Batts, 2008). Thus, range expansion of L. mazamae and/or L. cervi via movements of White-tailed Deer, albeit not rapid, is possible. Human-mediated dispersal (of host and parasites) represents another potential mechanism for bringing the ranges of the two species together. In most Virginia counties, except for areas in northwestern Virginia where chronic wasting disease has been reported, hunters harvesting deer are allowed to transport entire animals throughout the state (VDGIF, 2019-2020). As deer keds do not leave the host’s body for several hours after death of the animal, hunters transporting deer carcasses also are likely to be inadvertently transporting their associated parasites (Samuel & Trainer, 1972). DNA sequencing of L. cervi and its associated hosts has confirmed the presence of five bacterial pathogens (Anaplasma, Bartonella, Borrelia, Ehrlichia, and Rickettsia) and one protozoan parasite (7rypanosoma) in this species (Skvarla & Machtinger, 2019 and references therein). To date, L. mazamae has only been tested for and found to contain the Gram-negative bacterium Bartonella (Reeves et al., 2006; Souza et al. 2017). Transmission to hosts and spread of pathogens by deer keds is not yet fully understood for any of these disease agents (Skvarla & Machtinger, 2019). However, vertical transmission of Bartonella schoenbuchensis corrig. Dehio et al. to offspring has been recently confirmed in L. cervi, making it a potential vector for this pathogen (de Bruin et al., 2015). The close proximity of the currently known ranges of L. cervi and L. mazamae in Virginia suggests that physical overlap in this state and the surrounding areas is likely, thus setting up an interesting ecological situation. The extent and strength of any potential interactions between these two species, and the possibility for pathogen exchange between the two taxa, merit further investigation. ACKNOWLEDGEMENTS We thank Q. Davenport and A. Lantz for providing the deer specimens from which the deer keds were collected. We also thank L. Treado (VMNH) for the Z. mazamae images. Joe B. Keiper and two anonymous reviewers provided valuable comments that improved the manuscript. LITERATURE CITED Batts, G. K. 2008. An assessment of Quality Deer Management on a private hunt club in the Virginia Piedmont. MLS. thesis, Virginia Polytechnic Institute and State University, Blacksburg, VA. 111 pp. Baer-Lehman, M. L., T. Light, N. W. Fuller, K. D. Barry-Landis, C. M. Kindlin, & R. L. Stewart. 2012. Evidence for competition between /xodes scapularis and Dermacentor albipictus feeding concurrently on white- tailed deer. Experimental and Applied Acarology 58: 301-314. Bequaert, J. 1935. The American species of Lipoptena (Diptera, Hippoboscidae). Bulletin of the Brooklyn Entomological Society 30: 170. Bequaert, J. 1942. A monograph of the Melanophaginae, or ked-flies, of sheep, goats, deer and antelopes (Diptera, Hippoboscidae). Brooklyn Entomological Society, Brooklyn, NY. 220 pp. Bequaert, J. 1952. The Hippoboscidae or louse-flies (Diptera) of mammals and birds. Part I. Structure, physiology and _ natural history. Entomologica Americana 32: 1—209. Bequaert, J. 1954. The Hippoboscidae or louse-flies (Diptera) of mammals and birds. Part II. Taxonomy, evolution and revision of American genera and species. Entomologica Americana 34: 1—232. Bequaert, J. 1957. The Hippoboscidae or louse-flies (Diptera) of mammals and birds. Part Il. Taxonomy, evolution and revision of American genera and species. Entomologica Americana 36: 417-611. Carreno, R. A., L. A. Durden, D. R. Brooks, A. Abrams, & E. P. Hoberg. 2001. Parelaphostrongylus tenuis (Nematoda: Protostrongylidae) and other parasites of 82 BANISTERIA white-tailed deer (Odocoileus virginianus) in Costa Rica. Comparative Parasitology 68: 177-184. Collins, M., R. Knutti, J. Arblaster, J.-L. Dufresne, T. Fichefet, P. Friedlingstein, X. Gao, W. J. Gutowski, T. Johns, G. Krinner, M. Shongwe, C. Tebaldi, A. J. Weaver, & M. Wehner. 2013. Long-term climate change: Projections, commitments and irreversibility. Pp. 1029-1136 Jn T. F. Stocker, D. Qin, G.-K. Plattner, M. Tignor, S. K. Allen, J. Boschung, A. Nauels, Y. Xia, V. Bex, & P. M. Midgley (eds.), Climate Change 2013: The Physical Science Basis. Contribution of Working Group I to the Fifth Assessment Report of the Intergovernmental Panel on Climate Change. Cambridge University Press, Cambridge, United Kingdom, and New York, NY, USA. de Bruin, A., A. Docters van Leeuwen, S. Jahfari, W. Takken, M. Foldvari, L. Dremmel, H. Sprong, & G. Foldvari. 2015. Vertical transmission of Bartonella schoenbuchensis in Lipoptena cervi. Parasites & Vectors 8: 1-6. Deem, S. L., A. J. Noss, R. Villarroel, M. M. Uhart, & W. B. Karesh. 2004. Disease survey of free-ranging grey brocket deer (Mazama gouazoubira) in the Gran Chaco, Bolivia. Journal of Wildlife Diseases 40: 92-98. Dick, C. W. 2006. Checklist of world Hippoboscidae (Diptera: Hippoboscoidea). Field Museum of Natural History, Department of Zoology, Chicago. 8 pp. Farajollahi, A., W. J. Crans, D. Nickerson, P. Bryant, B. Wolf, A. Glaser, & T. G. Andreadis. 2005. Detection of West Nile virus RNA from the louse fly Icosta americana (Diptera: Hippoboscidae). Journal of the American Mosquito Control Association 21: 474-477. Genoways, H. H. & R. M. Timm. 2005. Mammals of the Cosigtina Peninsula of Nicaragua. Mastozoologia Neotropical 12: 153-179. Graciolli, G., C. A. Zucco, P. H. Cancado, & G. Mourao. 2011. Parasitism rates of Lipoptena guimaraesi and a new record of Lipoptena mazamae on Ozotoceros bezoarticus from the Central Pantanal wetlands in Brazil. Revista Brasileira de Parasitologia Veterinaria 20: 178-180. Halos, L., T. Jamal, R. Maillard, B. Girard, J. Guillot, B. Chomel, M. Vayssier-Taussat, & H. Boulouis. 2004. Role of Hippoboscidae flies as potential vectors of Bartonella spp. infecting wild and domestic ruminants. Applied and Environmental Microbiology 70: 6302— NO. 53, 2019 6305. Handley, C. O., Jr., & C. P. Patton. 1947. Wild Mammals of Virginia. Commonwealth of Virginia Commission of Game and Inland Fisheries, Richmond, VA. 220 pp. Kelsey, A., & J. Finch. 2018. Deer ked: a Lyme-carrying ectoparasite on the move. Cutis 102: 121—122. Kennedy, M. J., R. A. Newman, & G. A. Chalmers. 1987. First record of Lipoptena depressa (Diptera: Hippoboscidae) from Alberta, Canada. Journal of Wildlife Diseases 23: 506-507. Kern, W. J., Jr. 2003. Neotropical deer ked or Neotropical deer louse fly, Lipoptena mazamae Rondani (Insecta: Diptera: Hippoboscidae). Featured Creatures from the Entomology & Nematology Department, Florida Cooperative Extension Service EENY-308: 1-3. Linzey, D. W. 1998. The Mammals of Virginia. McDonald and Woodward Publishing Company, Blacksburg, VA. 459 pp. Liu, D., Y. Z. Wang, H. Zhang, Z. Q. Liu, H. Z. Wureli, S. W. Wang, C. C. Tu, & C. F. Chen. 2016. First report of Rickettsia raoultii and R. slovaca in Melophagus ovinus, the sheep ked. Parasites and Vectors 9: 600. Maa, T. C. 1965. A synopsis of the Lipopteninae (Diptera: Hippoboscidae). Journal of Medical Entomology 2: 233-248. Maa, T. C. 1969. A revised checklist and concise host index of Huippoboscidae (Diptera). Pacific Insects Monographs 20: 261-299. Maa, T. C., & B. V. Peterson. 1987. Hippoboscidae, Pp. 1271-1282 In J. F. McAlpine, B. V. Peterson, G. E. Shewell, H. J. Teskey, J. R. Vockeroth, & D. M. Wood (eds.), Manual of Nearctic Diptera, Vol. 2, Monograph 28. Agriculture Canada, Ottawa, ON, Canada. Moreira, R. F., L. D. C. Farezin, U. A. Souza, B. Z. D. Silva, D. B. Amorim, A. Girotto-Soares, L. E. Surita, M. M. Alievi, G. Graciolli, & J. F. Soares. 2019. Pupipara (Diptera, Hippoboscidae) in wild birds attended at a rehabilitation center in southern Brazil. Brazilian Journal of Veterinary Parasitology 28: 330- 332. Reeves, W. K., M. P. Nelder, K. D. Cobb, & G. A. Dasch. 2006. Bartonella spp. in deer keds, Lipoptena HIGHTOWER ET AL.: NEOTROPICAL DEER KED 83 mazamae (Diptera: Hippoboscidae), from Georgia and South Carolina, USA. Journal of Wildlife Diseases 42: 391-396. Romero-Castafion, S., B. G. Ferguson, D. Guiris, D. Gonzalez, S. Lopez, A. Paredes, & M. Weber. 2008. Comparative parasitology of wild and domestic ungulates in the Selva Lacandona, Chiapas, Mexico. Comparative Parasitology 75: 115-127. Samuel, W. M., & D. O. Trainer. 1972. Lipoptena mazamae Rondani, 1878 (Diptera: Hippoboscidae) on white-tailed deer in southern Texas. Journal of Medical Entomology 9: 104—106. Shock, B. C. 2014. Studying the natural history of the Florida panther: ectoparasites, piroplasms, and genetic diversity. Ph.D. dissertation, University of Georgia, Athens, GA. 140 pp. Skvarla, M. J., & E. T. Machtinger. 2019. Deer keds (Diptera: Hippoboscidae: Lipoptena and Neolipoptena) in the United States and Canada: New state and county records, pathogen records, and an illustrated key to species. Journal of Medical Entomology 56: 744-760. Small, R. W. 2005. A review of Melophagus ovinus (L.), the sheep ked. Veterinary Parasitology 130: 141-155. Souza, U., B. Dall’Agnol, T. Michel, A. Webster, G. Klafke, J. R. Martins, C. B. Kasper, T. C. Trigo, R. Ott, M. Maria de Assis Jardim, & J. Reck. 2017. Detection of Bartonella sp. in deer louse flies (Lipoptena mazamae) on gray brocket deer (Mazama gouazoubira) in the neotropics. Journal of Zoo and Wildlife Medicine 48: 532-535. VDGIF [Virginia Department of Game and Inland Fisheries]. 2019-2020. Hunting & Trapping in Virginia. Virginia Department of Game and Inland Fisheries, Henrico, VA. 72 pp. Westrom, D. R., & J. R. Anderson. 1992. The distribution and seasonal abundance of deer keds (Diptera: Hippoboscidae) on Columbian black-tailed deer (Odocoileus hemionus columbianus) in northern California. Bulletin of the Society of Vector Ecologists 17: 57-69. Banisteria, Number 53, pages 84—86 © 2019 Virginia Natural History Society Notes on the Parasitic Beaver Beetle, Platypsyllus castoris Ritsema, 1869 and Mouse Nest Beetle, Leptinus orientamericanus Peck, 1982 (Coleoptera: Leiodidae: Platypsyllinae) in Virginia Ralph P. Eckerlin Mathematics, Science and Engineering Division Northern Virginia Community College Annandale, Virginia 22003 reckerlin@nvcc.edu Arthur V. Evans Department of Recent Invertebrates Virginia Museum of Natural History 21 Starling Avenue Martinsville, Virginia 24112 arthurevans(@verizon.net ABSTRACT The parasitic Beaver Beetle (Platypsyllus castoris Ritsema, 1869) is documented from Virginia for the first time from specimens collected on American Beaver (Castor canadensis Kuhl, 1820) in Charles City, Fairfax, and Rappahannock counties. Another leiodid previously recorded from the Commonwealth, the Mouse Nest Beetle (Leptinus orientamericanus Peck, 1982), is reported from the Southern Red-backed Vole. Myodes gapperi (Vigors, 1830), a new Virginia host record. Keywords. Myodes gapperi, new host record, Southern Red-backed Vole. INTRODUCTION Leiodid beetles in the subfamily Platypsyllinae are dorsoventrally flattened and wingless mammalian ectoparasites that lack or have reduced eyes and are associated with rodents, shrews, and moles. The Beaver Beetle, Platypsyllus castoris Ritsema, 1869 (Fig. 1), is an ectoparasite of the American Beaver, Castor canadensis Kuhl, 1820, in North America and the Eurasian Beaver, Castor fiber Linnaeus, 1758, in Eurasia. Originally described as a flea, this species was later recognized as a beetle by LeConte (1872), who placed it in its own family Platypsyllidae, a taxon that was later considered a subfamily of the Leiodidae (Lawrence & Newton, 1982). These eyeless beetles have numerous rows of spines that permit their passage through beaver fur, but help to prevent the host from easily grooming them out. The American Beaver has a geographic range that extends from Alaska and Canada south of the Arctic Circle to Florida and northern Mexico (Helgen, 2005) and it is likely that the Beaver Beetle has a distribution similar to that of its host (Peck, 2006). The beaver ectoparasite was previously known in North America from seven provinces in Canada (AB, BC, NF, NS, ON, QC, SK) and 26 states in the United States (AK, AL, CA, FL, IA, ID, IL, LA, ME, MI, MN, MO, MS, MT, NC, ND, NE, NJ, NY, OK, OR, PA, SD, TX, WA, WY), the District of Columbia, as well as in Europe (Peck, & Newton, 2017). The specimens noted in this work are the first records of the species’ occurrence in Virginia. MATERIALS AND METHODS Three beavers killed by vehicles were examined for ectoparasites and their beetles given to Eckerlin who ECKERLIN & EVANS: PARASITIC BEETLES 85 Fig. 1. Platypsyllus castoris Ritsema from Charles City County, Virginia. (credit A.V. Evans) preserved them in 70% ethanol. Later, they were processed by overnight clearing in 10% potassium hydroxide solution, dehydrated in an ethanol series, cleared in xylene, and mounted to slides in Canada balsam. Leptinus beetles were similarly processed and mounted. Vouchers of both species have been deposited in the collections of the National Museum of Natural History (NMNH) and the Virginia Museum of Natural History (VMNH). Specimens collected on the George Washington Memorial Parkway (GWMP) have been deposited in the collections maintained at the GWNP, Turkey Run Park Headquarters, in McLean, Virginia. Three additional Virginia specimens of Platypsyllus castoris were collected and identified by Evans and retained in his collection (AVEC) in Richmond, Virginia, which is affiliated with the NMNH in Washington, D.C. All of these beetles were combed out of the fur just behind the shoulders of two beavers. One specimen was point-mounted, while the other two were preserved in 70% ethanol. RESULTS The five records of Beaver Beetles collected in Virginia bear the following locality data: Two females ex Castor canadensis, 19 January 1986, K.A. Strohmayer, 0.5km south of Belle Haven Marina, GWMP, Fairfax County, Virginia. Five males, two females ex C. canadensis, 19 February 1986, J. McBreen, Rt. 123, Lorton, Fairfax County, Virginia. Eight males, three females ex C. canadensis March 1986, S.W. Gotte, Lake Royal, Burke, Fairfax County, Virginia. One male ex C. canadensis, 4 May 2019, A.V. Evans and M.C. Odom, Harrison Lake National Fish Hatchery, 37.336728 N, 77.187224 W, Charles City County, Virginia. One male and one female ex C. canadensis, same collection data as previous record, except 29 August 2019, 37.342805'N, 77.188125 W. A search for additional Virginia records at the NMNH and VMNH yielded no additional specimens. However, holdings in the Virginia Tech Entomology Collection (VTEC) in Blacksburg, Virginia produced an additional record that included six Beaver Beetles collected in January 1992 by Tom Blount from a beaver in Shenandoah National Park, Rappahannock County, Virginia. These specimens are preserved in ethanol and were identified by Eric Day. Another ectoparasitic platypsylline, Leptinus orientamericanus. Peck, 1982, (Fig. 2) occurs in numerous localities in the United States east of the Mississippi River (AL, DC, DE, FL, GA, IL, IN, KY, MD, NC, NJ, NY, OH, PA, SC, TN, VA, WV) and has been recorded in association with seven species of small mammals in the orders Eulipotyphla (formerly Insectivora) and Rodentia (Peck & Newton, 2017). Hosts in Virginia are the Northern Short-tailed Shrew, Blarina brevicauda (Say, 1823), Star-nosed Mole, Condylura cristata (Linnaeus, 1758), and Eastern Mole, Scalopus aquaticus (Linnaeus, 1758) from the counties of Culpeper, Fairfax, Highland, King George, Montgomery, and Tazewell (Peck, 1982; Eckerlin & Painter, 1993). Eckerlin examined 113 Southern Red-backed Voles, Myodes gapperi (Vigors, 1830) in Virginia and found a single female L. orientamericanus on a Southern Red- backed Vole collected 2 April 1993, Flattop Mountain, Giles County, Virginia. This specimen, now deposited in the NMNH represents a new Virginia host and county record. Its association with this species is_ likely accidental simply because of the rarity of its occurrence. Additional specimens of L. orientamericanus housed in the VTEC were collected from the Virginia counties of Cumberland, Henry, and Patrick, all of which are new county distribution records. Fig. 2. Leptinus orientamericanus Peck (sex undetermined) from Montgomery County, Indiana. (credit K. Schnepp) 86 BANISTERIA ACKNOWLEDGMENTS We thank Steve Gotte, John McBreen, and Karl Strohmayer for providing Beaver Beetle specimens. Michael Odom of the United States Fish and Wildlife Service is acknowledged for his assistance in procuring the specimens of Platypsyllus castoris from Charles City County. Kal Ivanov (VMNH), Floyd Shockley (NMNH), Paul Marek and Jackson Means (VTEC) examined their respective collections for additional Virginia records. The Virginia Department of Game and Inland Fisheries kindly provided scientific collecting permits. Kyle Schnepp, Florida State Collection of Arthropods, Gainesville, FL, generously allowed us to use his image of Leptinus orientamericanus. Brent Steury offered useful suggestions that contributed to the overall quality of this note. LITERATURE CITED Eckerlin, R. P., & H. F. Painter. 1993. The star-nosed mole, Condylura cristata, a new host for Leptinus orientamericanus (Coleoptera: Leptinidae) in Virginia. Proceedings of the Entomological Society of Washington 95: 639. NO. 53, 2019 Helgen, K. M. 2005. Family Castoridae. Pp. 842-843 In D. E. Wilson & D. M. Reeder (eds.), Mammal Species of the World. 3 ed. Johns Hopkins University Press, Baltimore, MD. Lawrence, J. F., & A. F. Newton, Jr. 1982. Evolution and classification of beetles. Annual Review of Ecology and Systematics 13: 261-290. LeConte, J. L. 1872. On Platypsyllidae, a new family of Coleoptera. Proceedings of the Zoological Society of London 1872: 799-803. Peck, S. B. 1982. A review of the ectoparasitic Leptinus beetles of North America (Coleoptera: Leptinidae). Canadian Journal of Zoology 60: 1517-1527. Peck, S. B. 2006. Distribution and biology of the ectoparasitic beaver beetle Platypsyllus castoris Ritsema in North America (Coleoptera: Leiodidae: Platypsyllinae). Insecta Mundi 20: 85-94. Peck, S. B., & A. F. Newton. 2017. An annotated catalog of the Leiodidae (Coleoptera) of the Nearctic Region (Continental North America north of Mexico). Coleopterists Bulletin 71: 211-258. MISCELLANEA 87 Miscellanea Reports 1. Minutes of the Executive Committee of the Virginia Natural History Society Meeting of November 2, 2019. The 2019 meeting of the Executive Committee of the Virginia Natural History Society was called to order by President Nancy Moncrief at 9:04 a.m. on November 2, 2019 in the Board Room of the Virginia Museum of Natural History, Martinsville, Virginia. In attendance were Nancy Moncrief, Steve Roble, Kal Ivanov, Karen Powers, Paul Marek, Michael Lachance, Todd Fredericksen, Art Evans, and Curt Harden. The minutes were approved from the December 1, 2018 Executive Committee Meeting by unanimous consent. The revisions to Bylaws as edited on December 15, 2018 by Marek, which splits Secretary and Treasurer duties, allows concurrent terms for Councilors, and established Treasurer/co-Treasurer, were approved by unanimous consent. Announcements: Transfer of VNHS business office (funds and archive) from Hampden-Sydney to Virginia Museum of Natural History Foundation has been accomplished. Topics: Membership Report (Moncrief): As of October 23, there are nine student members, 58 regular members, and nine institutional members for a total of 76 members. Treasurer Report (Moncrief and Ivanov): On May 10, 2019, VNHS funds ($17,349.00) were transferred from Hampden-Sydney College to the VNHS business office. The costs of the 2019 Member Meeting were almost covered by the registration fees, and the difference will be paid to VMNH by the VNHS fund. These costs included nametags, poster boards and easels, lunches, and coffee breaks. Regarding the costs of back issues of Banisteria, Ivanov remarked that $15 per issue is prohibitively costly for students and early-career members. Roble stated that recently-retired faculty might be willing to donate their entire Banisteria library to interested parties. Other officers in attendance reminded the others that all Banisteria issues, excepting the current year, are free online. Ivanov remarked that $5 per issue (plus cost of shipping) is more reasonable, and if the member wants the entire collection of Banisteria, then it would be half off each issue. In addition, the member is obligated to pick up the collection in person from the VMNH archives. Nomination of officers was not required at this time, and was therefore not discussed. Date and location of the next joint Executive Committee and Member meetings: Evans mentioned that Randolph-Macon College (in Ashland, near Richmond) is a possibility. Evans argued that by being at a central location would leverage other members in_ the Commonwealth who might not be compelled to drive to the western part of the state. The Richmond area is good because there are many master naturalists in the area. However, Evans stated that parking is on the street and might be a concern. Randolph-Macon, as a possible location will be researched to determine room rental, lunch choices, street parking, and football scheduling. Moncrief, Powers, and Marek mentioned that the fall works well for their schedules. Powers and Marek stated that continuing the meetings on Saturday is compatible with teaching in the fall. A general discussion ensured about the necessity to avoid a Saturday when a football game at Randolph-Macon would be occurring. Powers mentioned that the Smithsonian center in Front Royal might be a good alternative. Finally, Moncrief mentioned that the 4-H center on Remount Road in Front Royal would be another prospect. (During the discussion with the members in the afternoon member Ellen stated that Charlottesville is yet a fourth possibility of a location for the meeting.) Moncrief remarked that the VMNH is a good location because the museum does not charge VNHS for room rental. Committee members present identified October 31, 2020 or November 7, 2020 (Saturdays) as appropriate choices of dates. These dates would necessitate a room for 9AM for the next Executive Committee Meeting. Following the Executive Committee meeting, a duration of 10am—4pm would allow 12-14 talks (Ivanov). Newsletter and Webmaster Report (Marek): Marek requested material for the next newsletter to be sent to him by December 1, 2019. Powers stated that master naturalist coordinator Michelle Prysby would be willing to send the newsletter to master naturalists. Marek will request a list of email addresses of current members from 88 BANISTERIA Moncrief and Ivanov to send the newsletter by January or February 2020. Moncrief stated that the next newsletter in 2020 might contain a section with “President’s Remarks”. Other items for the 2020 newsletter include a statement that Banisteria will be online-only starting in 2020. Moncrief provided revisions to the website to Marek at the Committee Meeting. A new website was necessitated and established for online payment by VNHS. Discussion ensured about transferring the old (.com) website to the new (.org) website with a different URL. Marek expressed concern regarding changing to a new URL since URL permanence in case published bibliographies refer to verbatim Banisteria URLs of papers would be broken after one or two years (this would occur even if an auto-redirect was established). Marek will contact Zach at the VMNH about the possibility of porting the payment widget from the new (.org) website to the preexisting (.com) VNHS website. Editors’ Report (Roble and Fredericksen): Roble remarked that he is working on obituaries for Banisteria #52. At 9:55AM, the Executive Committee Meeting was tabled until after the Members Conference. At 4:07PM, when the Executive Committee Meeting was reconvened to order, Roble remarked that issue #52 1s in the process of being finished, and will be printed and distributed by December 31, 2019. One hundred copies each of issues #52 and 53 will be printed, and they will be mailed together in the same envelope by Roble to all members who have paid dues 1n 2019 by the date Roble requests the members list from Moncrief and Ivanov. Issue #54 will be entirely online (2020). Marek will inquire with Virginia Tech Libraries regarding DOI generation for the first online issue. Evans recommended that issues 52 and 53 be printed together as a single 52-53 issue. Discussion ensued regarding this solution, and it was resolved that these last print issues will remain separate to demarcate the change of the editor from Roble to Fredericksen. Marek inquired if there would be one or two issues per year when Banisteria shifts to entirely online. Fredericksen stated that the published online-only articles each year would receive a DOI and be contained in a separate issue that would be dictated by the year (e.g., all articles published in 2020 will be in issue #54, all articles published in 2021 will be in issue #55, and so On). Other new business: Online repository of all the digital files including membership list will be placed on Google Drive. NO. 53, 2019 After the Conference, a Members’ meeting was called at 3:38 p.m. At the members’ meeting, Marek solicited feedback from the younger members of the society as to how the VNHS might serve their interests better. One member remarked that a Bioblitz might be held, which are beneficial for communicating to the public the importance of natural history and how scientists collect natural history data. A second member, suggested that bringing more master naturalists into the fold would be beneficial. This member stated that master naturalists like citizen science opportunities, as they can gain volunteer points towards their certification. The member additionally stated that these activities would also be helpful for undergraduates. A third member, recommend that we post the newsletter on the website, and this recommendation was seconded by Evans. A fourth member suggested that we have hands-on activities for the dual purpose of scientific outreach and to provide content for social media. A fifth member, suggested that these hands-on activities might include open labs and natural history collections, and soliciting natural history photos for an online gallery. The meeting was adjourned at 5:19 p.m. Respectfully submitted, Paul E. Marek, Secretary and Webmaster 2. President’s Report The second general meeting of the Virginia Natural History Society (VNHS) was held at the Virginia Museum of Natural History in Martinsville, VA on 2 November 2019. There were 40 attendees, 12 oral presentations, and 10 posters. I am pleased to report that 14 students (including two high school students) presented results of their research projects. The minutes of the VNHS Executive Committee meeting held on 2 November 2019 are included in this issue. The Committee approved the following changes to the VNHS bylaws: splitting Secretary and Treasurer duties, allowing concurrent terms for Councilors, and establishing Treasurer/co-Treasurer. The revised VNHS Bylaws are also included in this issue. The Committee also discussed holding the third general meeting of the Society at a location closer to Richmond. Subsequent to the meeting, we learned that 31 October 2020 will probably be the best date for the meeting. We are in communication with a potential venue and will announce the location and confirm the date of the 2020 meeting as soon as possible. MISCELLANEA 89 For a variety of financial and logistical reasons, Number 53 will be the last issue of Banisteria that will be printed and mailed. Future articles will be published online only. In addition, most or all of the Society’s business (announcements, ballots, dues notices) will transition to an electronic format. I encourage members to check the VNHS website frequently for announcements and to contact the VNHS Business Office to confirm that it has their current e-mail address. Respectfully submitted, Nancy D. Moncrief, VNHS President 3. Secretary-Treasurer’s Report On 1 May 2019, the VNHS business office was transferred from Hampden-Sydney College (HSC) to the Virginia Museum of Natural History (VMNH), and the VMNH Foundation assumed the role of fiscal agent for VNHS. As of 3 December 2019, total income for 2019 was $19,714.92, and total expenses were $2,019.50, leaving an account balance of $17,695.42. Details of income are as follows: 1) on 10 May 2019, a total of $17,349.61 was deposited from the account administered by HSC; 2) dues and contributions from May through 3 December totaled $1,815.31; 3) registration fees for the 2019 conference totaled $550.00. Details of expenses are as follows: 1) printing and mailing Banisteria 51 totaled $1,129.70; 2) website fees totaled $206.93; 3) nametags, easels, and posterboards for conferences totaled $184.97; 4) lunches and refreshments for the 2019 VNHS conference totaled $497.90. Issues 52 and 53 of Banisteria are scheduled to go to the printer by 31 December 2019, but those expenses will be paid in 2020. Respectfully submitted, Nancy Moncrief and Kal Ivanov, VNHS Co-Treasurers 4. Editor’s Report Banisteria is in a transition phase with a new editor and a new online publication format, starting in 2020 with volume 54. I thank the outgoing editor, Steve Roble, for his many years of service to the Society and his mentorship during the editorial transition. Online publication format of the journal will save printing costs, increase speed of distribution of the journal to members, and allow for color photographs without costs to authors. Alongside the changes in publication venue are proposed changes to the editorial structure. First, we would like to extend an invitation to a larger number (10-12) of Associate Editors (nominations are welcomed!), who would be consulted for reviewing or suggesting reviewers within their specialty areas of natural history. Also, a separate section of the journal will be reserved for papers involving citizen-science projects related to natural history. Possible contributors of papers to this section may include Virginia Master Naturalists, high school and college students, and members of nature centers and other non-government institutions. We hope that will increase inclusivity in the Society beyond teaching and research institutions and generate a broader audience to people with an interest in Virginia’s natural history. Please contact me if you have any comments or suggestions. Respectfully submitted, Todd Fredericksen Announcements 1. Dues renewal reminder Please renew your membership for 2020 by paying dues online or by sending a check payable to the Virginia Natural History Society to Co-Treasurer Dr. Nancy Moncrief. The website address for online payments 1s: https://virginianaturalhistorysociety.com/ membership/membership.htm You must provide an email address to receive future copies of Banisteria and other communications from the society. Student Member Incentive The Virginia Natural History Society is offering free one-year memberships for students (high school, undergraduate, or graduate) nominated by an advisor/ teacher/mentor who is a member in good standing of the Society. This offer is available for up to 20 students each year, and nominations will be considered in the order in which they are received. Nominators should include the following information for up to three students: name, institution, enrollment level, mailing address, e-mail address, and a short paragraph describing the student’s interests in and activities related to Virginia natural history. Nominations should be sent to the Chair of the membership committee, Dr. Kal Ivanov at kal.1vanov@vmnh. virginia. gov. 90 Article I. Article II. Article III. Article IV. Article V. Article I. BANISTERIA NO. 53, 2019 THE VIRGINIA NATURAL HISTORY SOCIETY ARTICLES OF INCORPORATION AND BYLAWS ARTICLES OF INCORPORATION NAME The society shall be called “The Virginia Natural History Society” (referred to hereinafter as “the Society’). OBJECTIVE The objective of the Society shall be to promote the study of all aspects of the natural history of Virginia, to educate the citizens of the Commonwealth about natural history, and to conserve its natural resources. The Society shall formulate bylaws to regulate its organization and procedures. The governing body of the Society shall be the Executive Committee. The Society is organized for scientific, educational, and charitable purposes as defined under the appropriate sections of the Internal Revenue Code. The Executive Committee is empowered to make appropriate changes to retain the tax-exempt status of the Society. Any such changes require a majority vote of the Executive Committee. BYLAWS MEMBERSHIP Section 1. Anyone who is interested in the objective of the Society is eligible for membership. Membership can be attained by application to the Secretary. Section 2. The classes of membership shall be Regular, Student, Family, Institutional, Supporting, Patron, Benefactor, Life, and Honorary. Student members must be currently enrolled as graduate, undergraduate, or high school students. A letter from their school verifying their student status must be included with the application for membership. Annual dues for these classes of membership shall be set by the Executive Committee and adjusted as needed. Honorary Councilors pay no dues. Section 3. Honorary Councilors shall be nominated by the Executive Committee and voted upon at the Executive Committee Meeting. Additional nominations may be made from the floor. Honorary Councilors shall be elected by a simple majority vote of the Executive Committee. Honorary Councilors shall be recognized leaders in the field of Virginia natural history. No more than 10 Honorary Councilors shall exist at any time. Honorary Councilors shall be eligible to hold office in the Society. Section 4. To be a Member in good standing a person must not be in arrears for dues. All annual dues for each ensuing year shall be due 1 January. Publications of the Society will be sent only to Members in good standing and to subscribing Institutions. Article II. MISCELLANEA 9] OFFICERS AND COMMITTEES Section 1. Section 2. Section 3. Section 4. Section 4.01 Section 4.02 Section 5. Section 6. Section 7. The officers of the Society shall be a President, a Vice-President (President Elect), a Secretary, a Treasurer, an Editor or co-editors, a webmaster, and three Councilors. The President shall be the chairman of the Executive Committee and presiding officer at meetings of the Society, and appoint any Standing Committees of the Society and any other committees deemed necessary. The President casts only tie-breaking votes. The President shall serve a 2-year term beginning January 1*t and ending December 31", but will usually not serve for two consecutive terms. The President shall make periodic or annual reports which are published in Banisteria. The Vice-President shall be the President-Elect, shall be elected by the plurality of ballots cast by the Society’s membership, and shall succeed to the Presidency in the event the office becomes vacant. The Vice-President shall be chairman of the Membership Committee, shall be responsible for coordinating with the local chairman arrangements for the scientific meetings of the Society, and shall preside over all meetings in the absence of the President. The Secretary shall be elected by a plurality of the ballots cast by the Society’s membership for a 4-year term. The Secretary shall record all the proceedings of the Society, supervise all official mailings including the Society’s publications, respond to all inquiries pertaining to membership, subscriptions, and Society matters, and maintain the membership and subscription lists. The Secretary shall compose a summary of the Committee and General Business Meetings. The Treasurer shall be elected by a plurality of the ballots cast by the Society’s membership for a 4-year term. The Treasurer shall be in charge of all funds, keep the financial records of the Society, and be responsible for an independent yearly audit. The Treasurer shall summarize the financial health of the Society and submit it for publication in Banisteria. The Secretary and Treasurer shall be responsible to the Executive Committee to whom an annual report shall be made. Editor or Co-editors shall be appointed to a 2-year term of office by the Executive Committee. Editor shall be responsible for preparation of the semiannual journal Banisteria, and for maintaining high scholarly standards in its content. The Editor shall be responsible to the Executive Committee to whom an annual report shall be made. The Webmaster shall maintain and update the Society website and provide an annual report to be published in Banisteria. The Executive Committee shall be the governing body and shall consist of the Executive Council in addition to the President, the Vice-President, the Secretary, the Treasurer, the Editor (or co-editors), Webmaster, and the last three Presidents. The Executive Council shall consist of three Councilors. Three Members in good standing who do not hold other Society offices shall be elected by a plurality of the ballots cast by the Society’s membership to serve 4-year terms as councilors. Councilors shall be eligible for reelection 2 years after the expiration of their previous terms. 92 Article III. Article IV. Section 8. Section 9. MEETINGS Section 1. Section 2. Section3. Section 4. Section 5 Section 6. ELECTIONS Section 1. Section 2. Section 3 Section 4 BANISTERIA NO. 53, 2019 A quorum of the Executive Committee shall consist of one more than half of its Members and must include the President or Vice-President. Decisions shall be made by simple majority of those Members present. Vacancies in the staff of officers shall be filled by appointments approved by the majority of the remaining Members of the Executive Committee, except in the case of presidential vacancy (see Article IH, Section 3). An appointee shall hold office only for the remainder of the term of his or her predecessor. Interim appointments of officers shall not constitute elected tenure. The Society shall sponsor such Scientific Meetings as it deems advisable. The objectives of the Society’s meetings shall be to present appropriate scientific papers and to foster the exchange of ideas among persons interested in Virginia natural history. There may be an annual General Meeting held sometime during a Scientific Meeting. Notice of this meeting shall be made with the notice of the Scientific Meeting (see Section 6 below). Those present shall constitute a quorum. The Executive Committee shall meet at some time during each meeting of the Society and at other times if deemed necessary by the President. As occasion demands, the Secretary, at the direction of the President, may submit matters to the Executive Committee for vote by mail or e-mail ballot. All such votes by the Executive Committee shall be placed on record and submitted for ratification at the next meeting of the Executive Committee. The Vice-President shall recommend the time and place of Annual Scientific Meetings. The Executive Committee shall give final approval to such recommendations. Notice of a Scientific Meeting shall be published in Banisteria, when possible. All Meetings, except Scientific Meetings, shall be conducted under Robert’s Rules of Order. The President, Vice-President, Secretary, Treasurer, and Councilors shall be elected by mail or on-line ballot from a list of nominees. Willingness of the nominees to serve shall be secured before their names are placed on the ballot. The Webmaster shall be appointed by the President. Ballots shall present a choice of at least two candidates for each of the offices of President-Elect, Secretary, Treasurer and Councilor if possible. The President shall notify the candidates for office of the election results and an announcement of the election results shall be published in the first issue of Banisteria following. A tie vote for any office shall be resolved by a secret ballot of the Executive Committee. Article V. Article VI. Article VU. MISCELLANEA 93 FINANCES Section 1. All funds received by the Society shall be used for publication of all official publications of the Society, and to defray other expenses incurred in the conduct of Society business as determined by the Executive Committee. Society Business includes but is not limited to the following: a. Publication of Banisteria. b. Publication of meeting programs. c. Rental fees for meeting rooms and monetary support of symposia, bioblitzes etc. d. Registration fees and provision of advanced monies for the purposes of scientific meetings. e. Annual audit. f. Postage and duplicating costs for ballots, reports for the Executive Meeting, and other official mailings of the Society. g. Maintenance of the website and domain. Section 2. No part of the net earnings of this Society shall ever inure to, or for the benefit of, or be distributable to its members, trustees, officers, or other private persons, except that the Society shall be empowered to pay reasonable compensation for services rendered, and to make payments and distributions in furtherance of the exempt purposes for which it was formed. Section 3. A yearly audit of Society finances shall be made, as provided in Article II, Section 4. Section 4. In the event that the Society shall cease to exist, after paying or adequately providing for the debts and obligations of the association, the remaining assets shall be distributed to a non-profit fund, foundation, or corporation, which 1s organized and operated exclusively for scientific, educational, and/or charitable purposes and which has established its tax exempt status under the appropriate Section of the Internal Revenue Code. The specific non-profit organization or organizations to receive any remaining funds may be determined and approved by the Executive Committee. PUBLICATIONS The Society shall publish such scientific publications as authorized by the Executive Committee. Members enjoy waived or reduced page charges when publishing in Banisteria. DUES Annual dues shall be determined by the Executive Committee and shall be due before the first day of each year. 94 Article VIII. Article IX. Article X. FISCAL YEAR BANISTERIA NO. 53, 2019 The fiscal year of the Society shall coincide with the calendar year. AMENDMENTS TO THE ARTICLES OF INCORPORATION Section 1. Section 2. Section 3. Proposed amendments to the Articles of Incorporation may be originated by the Executive Committee or by written request addressed to the Secretary and signed by at least 10 Members or by a simple majority of Members at a General Business Meeting of the Society. Voting on proposed amendments shall be by mail. The Secretary shall mail copies of proposed amendments and ballots to all members of the Society and Shall allow one month for their return, the due date being stated on the ballot. An affirmative vote by two-thirds of the ballots cast by the Society’s membership shall be necessary for adoption of an amendment to the Articles of Incorporation. Each proposed amendment shall be accompanied by a concise statement of its purpose, and comparison with the existing provisions, if any. The Secretary shall count and record the vote on an amendment and shall immediately notify the Executive Committee of the result. The result of the voting shall be announced to the membership of the Society in the next official issue of the journal. AMENDMENTS TO THE BYLAWS Bylaws for the conduct of the business of the Society may be enacted, amended, or repealed by a simple majority vote of the Executive Committee. Virginia Natural History Society http://virginianaturalhistorysociety.com/ General Information The Virginia Natural History Society (VNHS) was formed in 1992 to bring together persons interested in the natural history of the Commonwealth of Virginia. The VNHS defines natural history in a broad sense, from the study of plants, animals, and other organisms to the geology and ecology of the state, to the natural history of the native people who inhabit it. The goals of the VNHS are to promote research on the natural history of Virginia, educate the citizens of the Commonwealth on natural history topics, and to encourage the conservation of natural resources. Dissemination of natural history information occurs through publication of the society’s journal Banisteria and an annual newsletter. The first issue of Banisteria, named in honor of John Banister (1650-1692), the first university-trained naturalist to work in Virginia, was published in 1992. Articles cover a wide array of subjects, and prospective authors are encouraged to submit manuscripts on any aspect of natural history in Virginia; papers may pertain to Virginia or regional archaeology, anthropology, botany, ecology, zoology, paleontology, geology, geography, or climatology. Biographies, obituaries, and historical accounts of relevance to natural history in Virginia are also welcomed. Manuscripts are peer-reviewed for suitability and edited for inclusion in the journal. The society’s website contains detailed instructions for prospective authors and PDF reprints of all Banisteria articles that are more than two years old. Banisteria is indexed by Zoological Record and is available through EBSCO and the Biodiversity Heritage Library. Memberships The VNHS is open to anyone with an interest in natural history and welcomes participation by all members in society activities and efforts to promote education and conservation. Membership includes a subscription to Banisteria and invitation to an annual meeting. Annual dues for members are $20 (per calendar year); library subscriptions are $40 per year. Payments may be made online or via a check or money order sent to the Treasurer. Copies of most back issues of Banisteria are available for sale at a reduced price. The VNHS is a tax-exempt, nonprofit, society under Section 501(C)3 of the IRS. We welcome donations to support our mission in Virginia. Virginia Natural History Society Application for Membership Name Address Zip Code Phone Email Area(s) of Interest ANNUAL DUES AND SUBSCRIPTIONS TO BANISTERIA $500.00 Life (not annual) $300.00 Benefactor $100.00 Patron $50.00 Supporting $40.00 Institutional $25.00 Family $20.00 Regular $5.00 Student (see below) Contribution in addition to membership dues $ L L L L L eal L L L The special student rate is applicable only when accompanied by the following certification signed by a faculty advisor (students are also eligible for a 1-year free membership if an advisor’s nomination is approved by the society’s Executive Committee; see nomination guidelines in Banisteria). Institution Advisor Date Online membership registration and payments may be made at this website: https://www. virginianaturalhistorysociety.com/ membership/membership.htm If paying by mail, send membership form and dues (check or money order payable to Virginia Natural History Society) to: Dr. Nancy Moncrief, VNHS Treasurer Virginia Museum of Natural History 21 Starling Avenue Martinsville, VA 24112 gt Ci Disa S SUF ek Ranuncnlis NLL PL Frragartia tog .