Annals of the Entomological Society of America, 110(3), 2017, 269–275 doi: 10.1093/aesa/saw094 Advance Access Publication Date: 10 January 2017 Research Research article

Beetles (Insecta: Coleoptera) Associated With the , Geomys bursarius (Mammalia: Rodentia: Geomyidae), in Indiana

Gareth S. Powell,1,2 R. Michael Brattain,3 and Jennifer M. Zaspel1 Downloaded from https://academic.oup.com/aesa/article/110/3/269/2888452 by guest on 28 September 2021 1Department of Entomology, Purdue University, 901 West State St., West Lafayette, IN 47907 ([email protected]; jzaspel@purdue. edu), 2Corresponding author, [email protected], and 3505 Lingle Terrace, Lafayette, IN 47901 ([email protected])

Subject Editor: Michael Caterino

Received 23 March 2016; Editorial decision 21 November 2016

Abstract The beetle (Insecta: Coleoptera) fauna associated with the underground burrow systems of the plains pocket go- pher, Geomys bursarius (Shaw) (Mammalia: Rodentia: Geomyidae), is reported from one of Indiana’s primary conservation zones, Kankakee Sands, for the first time. Pitfall traps baited with pig dung were placed into active burrow systems and routinely checked from February 2014 to January 2015. A total of 26 species of Coleoptera were found in the burrow systems. We summarize phenological data for the most commonly collected species. Also, a complete checklist of burrow-inhabiting beetles is provided.

Key words: obligate, Scarabaeidae, Histeridae, Staphylinidae, inquiline

The Efroymson Restoration at Kankakee Sands is one of the largest within the underground burrow systems of the plains pocket gopher. prairie restorations east of the Mississippi River (7,800þ acres), lo- The burrows, which are kept isolated from the surface, resemble a cated on the far western boarder of northern Indiana (Nature cave system in which the gopher’s activities support a mostly closed Conservancy 2016). Historically, the areas in northern and ecosystem of organisms. The beetles and flies tend to inhabit the nest Indiana were composed of an estimated 400,000 to 1 million acres and fecal chambers while the cave crickets tend to inhabit the burrow of swamps, savannas, prairies, and forests known as the Grand runways (Thorne and Anderson 1990). In an effort to characterize the Kankakee Marsh. Beaver Lake, which covered a significant area in biodiversity associated with this system, insects were surveyed across what is currently known as Kankakee Sands Preserve, was drained an 11-mo period. Here, we document the beetle fauna living within a over several decades at the end of the 1800s. By 1917, the Kankakee sampling of burrows at the Kankakee Sands, Newton County, IN. River was ditched and straightened on the Indiana side, and the marsh itself was eliminated (McDowell et al. 1983, Nature Conservancy Materials and Methods 2016). Today, the Kankakee Sands site is commonly referred to as a “prairie peninsula,” located between the Kankakee and Iroquois River A population of plains pocket gophers was identified in an accessible basins. It is composed of large remnants of sand prairie, herbaceous restoration prairie in Newton County, IN. The specific field sites wetland, black oak barrens, and pine oak flat woods; these and other chosen were fire-managed several weeks before trapping was to be- natural areas are managed by the Indiana Department of Natural gin. This burning allowed the Geomys “push-ups,” which are small Resources (IDNR) and is an important conservation zone in the piles of sand created on the ground surface when a gopher is exca- Central Ecoregion (Nature Conservancy 2012). vating a fresh tunnel, to be more easily identified (Fig. 1A). These The hydric zones in this restoration range from emergent wetlands piles of sand bury plant life and then dry, first covering existing to small sand dunes (Nature Conservancy 2012). This area resides be- plant growth, leaving bare earth that can be discerned easily, but tween three prairie remnants, a strategy that was implemented to pro- eventually giving rise to new growth and in some cases even in- tect other areas from further fragmentation (Nature Conservancy creased growth. These piles eventually flatten back out and become 2012). The remaining prairie ecosystem is a small remnant of what it undiscernible amongst the tall grasses. The mounds show evidence once was, much having been grazed and cultivated by early settlers. of recent rainfall providing a visual gauge for the most recent under- Consequently, the Kankakee Sands restoration area provides habitat ground activity. A series of these fresh piles of sand in a row are in- for many uncommon and rare species for the area, including the plains dicative of a new tunnel or run being developed below the surface. pocket gopher, Geomys bursarius (Shaw). This species is geographi- As the year progressed, vegetation grew back in and the fresh “push- cally limited primarily to this area in Indiana (Quinn et al. 2010). A va- ups” became very hard to locate; it became imperative to adequately riety of insect inquilines (mainly beetles, cave crickets, and flies) live mark trap locations to allow for retrieval (Fig. 1B).

VC The Authors 2017. Published by Oxford University Press on behalf of Entomological Society of America. All rights reserved. For Permissions, please email: [email protected] 269 270 Annals of the Entomological Society of America, 2017, Vol. 110, No. 3 Downloaded from https://academic.oup.com/aesa/article/110/3/269/2888452 by guest on 28 September 2021

Fig. 1. Habitat sampled with Geomys push-ups present. (A) May 2014, (B) September 2014.

Methods for locating active burrows and removing the gopher With the vertical hole created bisecting an active tunnel, Victor follow Skelley (1992). A long, thin piece of metal, usually a 2-ft-long Easy-set Gopher Traps were set on either side, and then the vertical flathead screwdriver, was thrust into the ground between two re- shaft covered with a wooden board as to keep the subterranean cently created sand piles. A difference in resistance can be felt when qualities of the system intact. Rodent traps were checked the follow- the screwdriver breaks through the top of a tunnel and moves ing day; any gophers were collected, frozen, and then deposited in through the open area before hitting the more compact floor. A hole the Purdue University Vertebrate Collection. was dug straight down to the level of the tunnel floor, exerting con- Once the gopher was removed, a pitfall trap was placed in the siderable effort not to collapse the sides and keep the tunnel intact. center of the open portion of the tunnel. Plastic cups, 4 inches in Annals of the Entomological Society of America, 2017, Vol. 110, No. 3 271 Downloaded from https://academic.oup.com/aesa/article/110/3/269/2888452 by guest on 28 September 2021

Fig. 2. Depiction of baited pitfall trap placed into pocket gopher burrow system. Pitfall cup placed at burrow level, dung bait suspended above preservative, wooden cover board over hole and sealed with excavated sand. diameter, were filled part-way with ethylene glycol and placed with Suite version 4.2 software (Fig. 3). Data were compiled for each beetle the rim of the cup below the surrounding sand level. A small plastic species and phenology charts were created using the package “ggplot” container filled with moistened pig feces was suspended above the (Wickham, 2009) in RStudio v.0.99.879 (RStudio Team, 2016). Trap cup using wire (Fig. 2). These traps were serviced between a few catch data were averaged per day for each trapping period for each days and a few weeks after placement, depending on weather and species. These values were then pooled for each month, giving a single travel schedules. Shorter trapping periods were desired due to the value of total collected specimens of each species per month. This is vi- common behavior of the gophers to “back fill” disturbed tunnels. sualized for the nine most commonly collected species of beetle along This was observed most commonly in systems where the gopher was with the total Coleoptera catch (Fig. 4). not trapped out, but also occasionally in systems where one mam- Representatives of most beetle taxa, along with all nontarget ar- mal was removed. In these cases, it is possible that either another in- thropods, were deposited into the Purdue Entomology Research dividual was also active in that system, or, once the occupant was Collection (PERC), with select specimens covering most commonly removed, another found and colonized the system. found taxa retained by the authors. Representatives of the rove bee- Pitfall traps were active from February 2014 until January 2015. tles were deposited to the Field Museum of Natural History Trap placement varied based on recent gopher activity that was ob- (FMNH), and representatives of the scarab beetles were deposited in served. In general, traps were active at all times during the sampling the Florida State Collection of Arthropods (FSCA). period. However, the number of traps varied from six to ten due to availability of resources and current gopher activity. Collecting per- Results and Discussion mits issued for the removal of the mammals limited the number that were taken and also provided a period of time during the summer A total of 885 individual adult beetles were collected in total with where no gopher could be killed; this was done to limit the effect on baited pitfall traps (Table 1). Also collected, but not included in the the reproductive activity period of the gophers. These measures analysis, were several hundred true flies (Diptera) and camel crickets were taken so that the study as a whole did not impact the Geomys (Orthoptera: Rhaphidophoridae) (data not shown here). All population inhabiting the property. Coleoptera collected are presented here; however, it is likely many Occasionally pitfall traps were buried in sand but samples could species, namely, members of the families Carabidae, Monotomidae, be recovered using a floatation method. The preservative and sand Nitidulidae, and Phalacridae, occur only opportunistically in the mixture containing specimens was placed in a five-gallon bucket; a burrow systems and are not associated directly with the mammals. large quantity of water was then added to the mixture and agi- The raw trap data presented for each species broadly show the tated. With this method, organic debris floats to the surface and most commonly collected taxa; however, comparisons cannot be sand particles fall. The floating material was sieved and sorted un- considered comprehensive due to the limitations of the fieldwork. der a microscope. This process was repeated many times until no Traps were placed throughout the year as appropriate burrows were organic material was produced. All beetle specimens from the proj- found and gophers could be removed, but this led to variability in ect were curated and identified to lowest taxonomic level possible the trapping period length, number of pitfalls active on any given by the authors with expert input acquired for scarab and staphyli- day, and mammal presence in the burrow at the time of sampling. nid beetles. The number of pitfall traps ranged from six to ten in the ground at High-resolution dorsal habitus images were taken using a Leica any one time, and the duration each trap was active ranged from a DFC450 camera mounted onto a M165C stereomicroscope. few days to several weeks. However, any inconsistencies due to vari- Montaged dorsal habitus images were created using Leica Application able pitfall number were dispersed across the entire sampling period, 272 Annals of the Entomological Society of America, 2017, Vol. 110, No. 3 Downloaded from https://academic.oup.com/aesa/article/110/3/269/2888452 by guest on 28 September 2021

Fig. 3. Dorsal habitus images of a sampling of Coleoptera collected. (A) Dellacasiellus kirni (Cartwright). (B) Cryptoscatomaseter iowensis (Wickham). (C) Geomysaprinus rugosifrons (Wenzel). (D) Aleocharinae sp. 2. Annals of the Entomological Society of America, 2017, Vol. 110, No. 3 273 Downloaded from https://academic.oup.com/aesa/article/110/3/269/2888452 by guest on 28 September 2021

Fig. 4. Phenology charts for the nine most commonly captured species of Coleoptera and total Coleoptera across the study. Collection frequency is presented on the y-axis, numbers were pooled for each month.

as patterns causing those inconsistencies were constant. Also, pitfall collected species (i.e., Aleocharine sp. 1 and sp. 2, traps placed in systems during the time where the gopher was not to Cryptoscatomaseter iowensis, and Geomyphilus insolitus) depicted be removed were not under grossly different conditions, as gopher a bimodal distribution, with both spring and fall activity periods. In activity within a system often continued after the removal of a single general, the total Coleoptera catches followed the same pattern, specimen. Despite these possible limitations, the resulting raw with both spring and fall periods of increased activity with greatly frequencies show some clear patterns (Fig. 4). For example, some reduced to absent faunas recovered the rest of the year. species (i.e., Atholus minutus, Cryptoscatomaseter punctissimus, Comparisons between the results presented here and previous Dellacasiellus kirni, and Geomysaprinus rugosifrons) each have sin- survey work done with gopher burrow inhabitants across North gle peaks of adult activity, with A. minutus, D. kirni, and G. rugosi- America are presented in Table 2. Not all surveys provided exhaus- frons clearly active in the spring, and C. punctissimus almost tive data for all families of beetles. However, some broad similarities exclusively collected in November. In others, very few specimens along with some contrasts between results can be discussed. Both were found throughout the year, but specific instances where adults the study that took place in Wisconsin (Kriska and Katovich 2005) were up to 50 times more abundant emerged from our dataset show- and another in (Paulsen 2006) only present species of the ing periods of increased subterranean adult activity (i.e., Atholus family Scarabaeidae. Another variable of note was summarized by minutus, Chilothorax distinctus). The remaining commonly Tishechkin and Cline (2008), the species of mammal present 274 Annals of the Entomological Society of America, 2017, Vol. 110, No. 3 through these studies differs by location: Geomys breviceps (Baird) and Texas (Peck and Skelley 2001); from these studies in compari- in Louisiana and Texas; G. bursarius in Arkansas, Indiana, son with ours, obligate pocket-gopher inhabiting leiodids may be re- Nebraska, and Wisconsin; and G. pinetis Rafinesque in Florida. stricted to the southern United States. Identical diversities of scarab Tishechkin and Cline (2008) found 22 obligate beetle species in go- beetles were found in both the present study and Tishechkin and pher systems across Louisiana; this general species diversity is also Cline (2008), with several species being consistent across the two found in our study (i.e., 18 beetle species). Some key differences in- studies; however, all scarabs sampled in Indiana were aphodiines clude the presence of a fourth family, Leiodidae, in the Louisiana whereas in Louisiana they also collected a species of Euphoria study. This family was also discovered in similar sampling in Florida (Scarabaeidae: Cetoniinae). Comparisons of the staphylinid fauna between the studies are limited at this time due to the presence of one confirmed undescribed species along with several suspected new Table 1. Beetle taxa collected by subterranean pitfall traps with to- taxa of Aleocharinae in our sampling. Until further taxonomic stud- tal abundance data ies can be conducted, the composition of this beetle family in gopher List of all Coleoptera species found in burrows with total abundance systems in Indiana is not well understood. Similarities were also Downloaded from https://academic.oup.com/aesa/article/110/3/269/2888452 by guest on 28 September 2021 data found with the Histeridae, Atholus minutus was collected in both IN and LA, and members of the genus Geomysaprinus were also Carabidae present in both regions. Amara sp. 1 Several other rigorous surveys of the obligate fauna associated Harpalus erraticus Say 1 Trechus sp. 2 with pocket gophers have been undertaken in recent years (Skelley Cryptophagidae and Woodruff 1991, Peck and Skelley 2001, Skelley and Gordon Henoticus sp. 1 2001). As with Tishechkin and Cline (2008), broad family-level sim- Histeridae ilarities were also present between studies, but individual species Atholus minutus (Ross) 14 varied from one location to another. Recently, Kovarik et al. (2008) Geomysaprinus rugosifrons (Fall) 66 summarized the obligate beetle fauna for pocket gophers in Hypocaccus sp. nov. 1 Arkansas. This study identified five species of Scarabaeidae and five Monotomidae species of Histeridae; several of these were the same taxa collected Europs pallipennis (LeConte) 1 at Kankakee Sands (A. minutus, D. kirni, G. insolitus). The absence Nitidulidae Glischrochilus quadrisignatus (Say) 2 of Leiodidae in Kovarik et al. (2008) provides additional evidence to Cryptarcha ampla Erichson 1 the distribution of obligate species being exclusively southern in Phalacridae distribution. Staphylinidae collected in the Arkansas study were Stilbus sp. 2 excluded, so cannot be compared but were certainly in the systems Staphylinidae and much like the specimens from Indiana, require further study. Aleocharine sp. 1 113 Although previous formal surveys of the burrow-dwelling Aleocharine sp. 2 228 Coleoptera fauna in the pocket gopher systems have been under- Anotylus sp. 7 taken, here we present a dataset for Indiana for the first time. With Bledius sp. 1 this preliminary study completed, a baseline exists for future surveys Quedius sp. nov. (male) 1 in the Ohio River valley. The range for the plains pocket gopher Quedius sp. (female) 2 Sepedophilus sp. 1 extends west throughout the Great Plains, where more faunistic re- Scarabaeidae search is needed. Much of the range of Geomys remains under- Ataenius abditus (Haldeman) 5 studied, with the obligate fauna not known for many populations. Chilothorax distinctus (Muller) 25 There is high potential for isolation with burrow-inhabiting organ- Cryptoscatomaseter iowensis (Wickham) 220 isms, where systems are commonly not connected and where the Cryptoscatomaseter magnificens (Robinson) 1 host populations are also often fragmented. Further studies could Cryptoscatomaseter punctissimus (Brown) 49 lead to new taxa as well as a better understanding of the current ob- Dellacasiellus kirni (Cartwright) 93 ligate fauna. This study also provides the Kankakee Sands Geomyphilus insolitus (Brown) 44 Restoration with preliminary phenology data for Coleoptera associ- Platytomus atlanticus (Cartwright) 3 Total 885 ated with a mammal species extremely restricted geographically for the state.

Table 2. Comparison of beetle faunas of Geomys burrows across multiple studies (in part adapted from Tishechkin and Cline 2008, with additional data included from Kovarik et al. 2008)

Number of species

Family of Coleoptera Indiana Arkansas Florida Louisiana Nebraska Texas Wisconsin

Histeridae 3 (2) 5 7 4 No Data 5 No Data Leiodidae 0 No Data 8 2 No Data 4 No Data Scarabaeidae 8 (6) 5 12 7 13 9 6 Staphylinidae 7 (5) No Data No Data 8 No Data No Data No Data

Studies summarized here represent several different mammal species—Geomys breviceps (LA, TX), G. bursarius (AR, IN, NE, WI), G. pinetis (AL, FL, GA). Data given for this study have total species per family with the number of presumed obligates in parentheses. Annals of the Entomological Society of America, 2017, Vol. 110, No. 3 275

Acknowledgments Nature Conservancy. 2012. Internal Site Conservation Plan for the Kankakee Sands Macrosite. (internal conservation and business plan). We thank Ted Anchor and John Shuey (Nature Conservancy) for access to the Nature Conservancy. 2016. Illinois-Indiana, Kankakee Sands: A brief history. property, permission to collect insect specimens, and general support of the (http://www.nature.org/ourinitiatives/regions/northamerica/unitedstates/ project. Also, Roger Hedge and Linnea Petercheff (Indiana Department of indiana/placesweprotect/kankakee-sands-history.xml) Natural Resources) are thanked for providing the necessary Scientific Paulsen, M. J. 2006. A new species and new records of Aphodius Illiger Purposes License for mammal trapping. Alfred Newton and Margaret Thayer (Coleoptera: Scarabaeidae: Aphodiinae) from mammal burrows in (FMNH) assisted with identification of staphylinid specimens, and Paul Nebraska. Insecta Mundi 20: 95–100. Skelley (FSCA) confirmed determinations for several scarab beetles. Gino Peck, S. B., and P. E. Skelley. 2001. Small carrion beetles (Coleoptera: Nearns (PERC), Crystal Maier (FMNH), and Paul Skelley are thanked for Leiodidae: Cholevinae) from burrows of Geomys and Thomomys pocket aiding in the deposition of vouchers in the mentioned institutions. Kyle gophers (Rodentia: Geomyidae) in the United States. Insecta Mundi 15: Schnepp (FSCA) is sincerely thanked for performing preliminary fieldwork 139–149. that made this study possible. Jeff Holland and Adam Alford provided assis- Quinn, V. S., C. C. Tsai, and P. A. Zollner. 2010. Distribution of the Plains tance with analysis and summary of data. Prairie Biotic Research Inc. is gra- Pocket Gopher (Geomys bursarius) in the grassland physiographic regions ciously acknowledged for partially funding this project. Paul Skelley and one of Indiana. Proc. Indiana Acad. Sci. 119: 87–94. Downloaded from https://academic.oup.com/aesa/article/110/3/269/2888452 by guest on 28 September 2021 anonymous reviewer, along with editor Michael Caterino, are sincerely Ross, E. S. 1940. New Histeridae (Coleoptera) from the burrows of the thanked for comments that improved this manuscript. Florida pocket gopher. Ann. Entomol. Soc. Am. 33: 1–9. RStudio Team 2016. RStudio: Integrated Development for R. RStudio, Inc., References Cited Boston, MA. (http://www.rstudio.com) (Accessed April 2016). Skelley, P. E. 1992. Trapping Pocket Gopher Insects. Scarabs. No. 6, 4. (http:// Connior, M. B. 2011. Geomys bursarius (Rodentia: Geomyidae). Mammalian museum.unl.edu/research/entomology/Newsletter/scarabs_6.pdf) Species 43: 104–117. 879 Skelley, P. E., and R. D. Gordon. 2001. Scarab beetles from pocket gopher Gordon, R. D., and P. E. Skelley. 2007. A monograph of the Aphodiini in- burrows in the southeastern United States (Coleoptera: Scarabaeidae). habiting the United States and Canada (Coleoptera: Scarabaeidae: Insecta Mundi 15: 77–93. Aphodiinae). Mem. Am. Entomol. Inst. 79: 1–580. Skelley, P. E., and R. E. Woodruff. 1991. Five new species of Aphodius Kovarik, P., S. Chordas, III, H. Robinson, P. Skelley, M. Connior, J. Fiene, (Coleoptera: Scarabaeidae) from Florida pocket gopher burrows. Fla. and G. Heidt. 2008. Insects inhabiting the burrows of the Ozark pocket go- Entomol. 64: 517–536. pher in Arkansas. J. Ark. Acad. Sci. 62: 75–78. Thorne, D. H., and D. C. Anderson. 1990. Long-term soil-disturbance pattern Kriska, N., and K. Katovich. 2005. Scarab beetles (Coleoptera: Scarabaeidae) by a pocket gopher, Geomys bursarius. J. Mammol. 71: 84–89. associated with pocket gophers in Wisconsin. Great Lakes Entomol. 38: Tishechkin, A. K., and A. R. Cline. 2008. The beetle (Coleoptera) fauna of 2–50. pocket gopher burrows in Louisiana. Proc. Entomol. Soc. Wash. 110: McDowell, B., J. Newman, and J. Ebinger. 1983. Survey of the woody vegeta- 331–339. tion of the Kankakee Sand Area Section of Indiana and Illinois. Proc. Wickham, H. 2009. ggplot2: Elegant Graphics for Data Analysis. Springer- Indiana Acad. Sci. 93: 187–194. Verlag, New York.