Dung Beetle (Coleoptera: Scarabaeidae and Geotrupidae) Communities of Eastern Maryland Author(S) :Dana L

Dung Beetle (Coleoptera: Scarabaeidae and Geotrupidae) Communities of Eastern Maryland Author(s) :Dana L. Price, Lauren M. Brenneman and Ryan E. Johnston Source: Proceedings of the Entomological Society of Washington, 114(1):142-151. 2012. Published By: Entomological Society of Washington DOI: http://dx.doi.org/10.4289/0013-8797.114.1.142 URL: http://www.bioone.org/doi/full/10.4289/0013-8797.114.1.142

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DUNG BEETLE (COLEOPTERA: SCARABAEIDAE AND GEOTRUPIDAE) COMMUNITIES OF EASTERN MARYLAND

DANA L. PRICE,LAUREN M. BRENNEMAN, AND RYAN E. JOHNSTON

Department of Biological Sciences, Salisbury University, 1101 Camden Avenue, Salisbury, Maryland 21801, U.S.A. (e-mail DLP: [email protected])

Abstract.—We examined the species diversity and seasonal abundance of 19 species of dung beetles (Coleoptera: Scarabaeidae and Geotrupidae) in two ecotone habitats of Maryland’s Eastern Shore. Specimens were collected over an eight-month period during 2009–2010 using standard pitfall trap transects of 200 m. In 2010, an additional survey of eight habitats in Wicomico and Worcester counties was conducted to obtain a more representative sample of the original dung beetle diversity. Habitat data, bait attraction, and collection methods are provided. This research is in- tended to be the ﬁrst bioinventory of Maryland dung beetles for conservation research. Key Words: bioinventory, Nassawango Creek Preserve, Pocomoke State Forest, seasonal abundance, species diversity, United States DOI: 10.4289/0013-8797.114.1.142

Dung beetle feeding and nesting be- conducted to sample dung beetles from haviors play a critical role in nutrient re- the Mid-Atlantic region of the United cycling, soil structure, and the removal States (Price 2004, 2006). The current of dung in both agricultural and natural knowledge of Maryland dung beetles is habitats. They are especially important limited to a checklist of Scarabaeoidea for consuming and burying dung that based on museum and literature review may otherwise create suitable breeding (Staines 1984), with limited reports from habitats for parasitic nematodes and flies counties in eastern Maryland. This re- (Nichols et al. 2008). Dung beetles are gion of Maryland is part of the Mid- also among the most cost-effective of all Atlantic Coastal Plain that extends from animal taxa for assessing and monitoring south of Long Island to the South Atlantic biodiversity (Gardner et al. 2008), and Coastal Plain at the Virginia-North Carolina consequently they are considered a model border. Pine forests dominate the outer group for understanding broad biodiver- Coastal Plain (nearer the coast), and hard- sity trends (Spector 2006). wood forests dominate the inner Coastal Long-term studies of any group of Plain. The boundary between the Atlantic Scarabaeinae are limited (Howden and Coastal Plain and the Piedmont of western Howden 2001). Few studies have been Maryland is known as the Fall Line, a transitional zone where harder rock of the Piedmont comes in contact with * Edited by Jens Prena; accepted by Robert R. sand, silt, and clay of the coastal sedi- Kula ments (Hanner et al. 2001). VOLUME 114, NUMBER 1 143

Our research objectives were to deter- water and salt. The bait was wrapped in mine the species diversity and seasonal cheesecloth, tied with a piece of biode- abundance of dung beetles (Coleoptera: gradable twine, and suspended from hard- Scarabaeidae and Geotrupidae) in two ware cloth inside the container. Plywood ecotone habitats of eastern Maryland, tiles were nailed into the ground at a 45 as well as to document natural history degree angle above the container to pre- data for species collected. The knowledge vent rainwater flooding. At each site, of species composition of an area can beetles were collected every seven days provide valuable information for moni- from June–October 2009 and March–June toring introduced species and provide a 2010, and traps were re-baited each time. baseline of the species composition within To determine habitat preference of the region. species collected during the initial study, and to collect beetles not attracted to MATERIALS AND METHODS dung or which are perhaps rare, further Dung beetles were sampled from June– sampling was conducted in Wicomico October 2009 and March–June 2010 in and Worcester counties from March– two ecotone habitats of Maryland’s East- December 2010. Six pitfall traps (de- ern Shore: (1) Nassawango Creek Pre- scription as above) baited with dung, carrion, fruit, and sometimes malt were serve (NCP: Wicomico County) and (2) set in four locations (two sites per loca- Pocomoke State Forest (PSF: Worcester tion, three traps per site). Site descriptions County). Ecotone habitats were chosen are listed below in Table 1. In addition to for this study to increase the diversity of our extended sampling efforts, The De- beetles inventoried (Spector 2002). In the partment of Natural Resources (DNR) NCP site, the traps were placed along donated 155 dung beetle specimens that a power line cut on the edge of the forest. were captured in a two-year study to The NCP consists of 50–80 year old survey the ant species diversity of dune mixed pine and hardwood forest that has habitats of Worcester County. Specimens been a conservation priority for The Nature were captured in unbaited pitfall traps Conservancy since 1978, with more than from 27 different sites. Additional sam- 9,000 acres now in Conservancy owner- pling methods included hand collection ship (The Nature Conservancy 2009). In from different sites, when possible, and the PSF site, traps were set along a dirt the use of a blacklight. road between a shortleaf pine forest and Aphodiinae specimens were identi- ; a plot of small pines ( 5–8 years old) that fied by Paul Skelley of the Florida State followed a recent harvest for Christmas Collection of Arthropods. Trogidae spec- trees. The PSF region consists of nearly imens were identified by Robert Androw 15,000 acres of loblolly pine, mixed pine- of the Carnegie Museum of Natural His- hardwood, shortleaf pine, and bald-cypress tory. Vouchers are housed in the personal forests that abut the Nassawango Creek. collection of Dana L. Price at Salisbury Using standardized pitfall trap tran- University in Salisbury, Maryland. These sects, five traps were baited with human specimens are serving as reference col- dung and placed 50 m apart along a lin- lection for future research. ear transect (200 m) at each site (Larsen We used EstimateS 8.20 (Colwell 2009) and Forsyth 2005). Traps consisted of to examine all data from the NCP and PSF. a 2.5 qt plastic container buried to the Samples input into EstimateS were based rim and filled with a small amount of on weekly captures (33 total samples for 144 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Table 1. Description of eight sites used for the collection of dung beetles (Scarabaeidae and Geotrupidae) in Wicomico and Worcester counties.

County Site Description

Wicomico 1 Open field (Powerline cut) in Nassawango Creek Preserve. This area was mowed once during 2010. 2 Mixed hardwood forest that runs parallel to the powerline cut. 3 Mixed pine forest located in the Nassawango Creek Preserve at the Nassawango Field Station (owned by Salisbury University). 4 Mixed pine and hardwood forest adjacent to location described above (Site 3). 5 Hardwood forest located at the Pemberton Historical Park. Pemberton, situated along the Wicomico River, consists of upland pines, hardwood forests, and wetland habitats. 6 Open unmowed field situated in Pemberton Historical Park (see above description) Worcester 7 Field of five-eight year old pines following a recent harvest. Located in the Pocomoke State Forest (PSF). 8 Shortleaf Pine forest. Located in the PSF. DNR Twenty-seven different sites located within the Chesapeake State Forest, Samples Nassawango Creek Preserve, Pocomoke State Forest, and Shad Landing State Park. each site). To estimate species richness is considered one of the most meaningful we used Chao1, (Chao 1984, Colwell diversity measures because it captures and Coddington 1994), ICE (Incidence- the variance of the species abundance base Coverage Estimator), and ACE distribution. (Abundance-base Coverage Estimator) (Chazdon et al. 1998, Chao et al. 2000). RESULTS AND DISCUSSION Species accumulation curves were used A total of 19 species (5,513 individuals) to illustrate the rate at which new species were collected from the Nassawango were sampled (Magurran 2004). Creek Preserve (NCP) and 17 species We used a rank/abundance plot to vi- (6,062 individuals) from the Pocomoke sualize the species abundance distribution State Forest (PSF) during the first year of of samples collected from 2009–2010 sampling (Table 2). Only six species (Table 2). In this, species are plotted made up 88% of the total individuals in sequence from most to least abun- collected. In the NCP location, with dant (Magurran 2004). To examine mixed hardwoods and pine, Onthopha- species diversity we used several non- gus pennsylvanicus Harold and Copris parametric diversity measures, including minutis (Drury) were the most abundant the Shannon index (H), Shannon ex- species with 1,066 and 1,023 individuals ponential mean (eH), and Simpson (in- verse) diversity index (1/D). Though collected, respectively. In the PSF loca- frequently reported, the Shannon in- tion, with sandy soil and mixed-pine dex often makes interpretation difﬁcult forests, Onthophagus tuberculifrons Harold (Magurran 2004); thus, more recent and Canthon pilularis (Linnaeus) were studies have used Shannon exponen- the most abundant species with 1,750 tial mean, which gives the number of and 1,221 individuals collected, respec- species that are found in the sample tively. This is congruent with Staines had all species been equally common (1984) that O. tuberculifrons is more (Whittaker 1972). The Simpson’s index commoninsandysoils. VOLUME 114, NUMBER 1 145

Table 2. Species list and abundance of scarabaeoid dung beetles (Coleoptera: Scarabaeidae and Geotrupidae) present in two locations of eastern Maryland. Species richness, abundance, and diversity at each site are shown at the bottom.

Species Name Nassawango Creek Preserve Pocomoke State Forest

Ataenius strigatus (Say) 1 0 Ateuchus lecontei Harold 1 6 Blackburneus aegrotus (Horn) 436 122 Blackburneus rubeolus (Palisot de Beauvois) 20 62 Canthon pilularius (Linnaeus) 755 1221 Copris minutis (Drury) 1023 814 Geotrupes. blackburnii (Fabricius) 192 233 Geotrupes egeriei Germar 534 875 Labarrus pseudolividus (Balthasar) 3 1 Melanocanthon bispinatus (Robinson) 12 100 Onthophagus concinnus Laporte 6 57 Onthophagus hecate (Panzer) 611 462 Onthophagus nuchicornis (Linnaeus) 1 0 Onthophagus pennsylvanicus Harold 1066 306 Onthophagus striatulus (Palisot de Beauvois) 8 3 Onthophagus taurus (Schreber) 3 7 Onthophagus tuberculifrons Harold 616 1750 Oscarinus rusicola (Melsheimer) 5 2 Phanaeus vindex MacLeay 220 16 Species Richness 19 17 Total Abundance 5513 6062 Estimated Species Richness Chao1 22 17 ICE 21 17.34 ACE 21 17.53 Species Diversity Shannon index (H) 2.13 1.98 Shannon exponential mean (eH) 8.44 7.27 Simpson’s index (1/D) 7.48 5.74

Estimated species richness was higher Seasonal trends were apparent in both overall for the NCP location with sites (Table 3). In the NCP, O. pennsyl- a Chao1 of 22, an ICE of 21, and an ACE vanicus was most abundant in May and of 21 (Table 2). Species accumulation C. pilularis in June (Fig. 3). Species curves are congruent with these ﬁndings that exhibited two generations per year indicating another two to three more were C minutus, abundant in March and species may be found in this location September–October, Onthophagus hecate (Fig. 1). Rank abundance curves suggest (Panzer), abundant in April and September, that although the NCP site has higher and O. tuberculifrons, abundant in May species richness, both sites are similar in and September–October. In the PSF, their evenness (Fig. 2). Overall the NCP C. pilularis was abundant throughout site had the highest diversity with a the summer from May until August, Shannon index of 2.13, a Shannon ex- O. tuberculifrons was most abundant dur- ponential mean of 8.44, and a Simpson ing May and September, Geotrupes egeriei index of 7.48 (Table 2). Germar during August, and C. minutis 146 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Fig. 1. Species accumulation curves for dung beetles sampled in the Nassawango Creek Preserve (NCP) and Pocomoke State Forest (PSF). The NCP site is shown in black, and the PSF site is shown in gray.

August–October (Fig. 4). It would have additional baits yielded 18 species and been interesting to continue collections 1,322 individuals, with just two new throughout the entire year to document species found: Chilothorax distinctus the decline of C. minutis and O. tuber- (Mu¨ller) and Dichotomius carolinus culifrons in the PSF habitats. Potentially (Linnaeus) (Table 4). We believe that we might have collected a few Apho- Chilothorax may have been overlooked diinae that are winter specialists. during the ﬁrst study, and only one speci- Collections of scarabaeoid dung bee- men of Dichotomius was collected, sug- tles during March–December 2010 using gesting that it may be rare at these localities.

Fig. 2. Rank abundance curve for the dung beetle communities indicated in Table 2. The Nassa- wango Creek Preserve is shown in black, and the Pocomoke State Forest is shown in gray. OUE14 UBR1 NUMBER 114, VOLUME

Table 3. Seasonal diversity and abundance of dung beetles (Coleoptera: Scarabaeidae and Geotrupidae) collected in the Nassawango Creek Preserve and Pocomoke State Forest.

March April May June July August September October Species Name NCP PSF NCP PSF NCP PSF NCP PSF NCP PSF NCP PSF NCP PSF NCP PSF Total

Onthophagus tuberculifrons Harold 7 13 80 227 164 429 11 56 3 48 10 18 153 771 188 188 2366 Canthon pilularius (Linnaeus) 0 0 1 5 92 403 335 238 154 308 80 148 90 118 3 1 1976 Copris minutis (Drury) 220 45 62 36 42 30 0 4 17 11 97 216 233 231 352 241 1837 Geotrupes egeriei Germar 10 2 2 8 1 35 13 15 25 110 105 463 253 133 125 109 1409 Onthophagus pennsylvanicus Harold 0 0 0 4 491 111 164 102 176 38 177 32 58 19 0 0 1372 Onthophagus hecate (Panzer) 0 2 192 66 128 102 61 60 124 106 62 121 26 25 18 5 1098 Blackburneus aegrotus (Palisot de Beauvois) 0 0 0 0 1 0 194 7 63 10 39 31 135 74 4 0 558 Geotrupes blackburnii (Fabricius) 88 61 17 30 0 10 0 2 0 0 0 0 18 0 69 130 425 Phanaeus vindex MacLeay 0 0 1 1 25 6 38 1 65 1 37 4 51 3 3 0 236 Melanocanthon bispinatus (Robinson) 0 0 0 0 0 29 2 63 6 2 4 5 0 1 0 0 112 Blackburneus rubeolus (Horn) 0 0 0 0 17 61 2 1 1 0 0 0 0 0 0 0 82 Onthophagus concinnus Laporte 0 0 3 23 1 28 0 2 1 3 0 1 1 0 0 0 63 Onthophagus striatulus (Palisot de Beauvois) 0 0 0 0 0 0 3 1 0 1 4 1 1 0 0 0 11 Onthophagus taurus (Schreber) 0 0 0 0 0 0 0 1 2 4 0 0 1 2 0 0 10 Ateuchus lecontei Harold 0 0 0 0 0 2 0 2 0 0 1 2 0 0 0 0 7 Oscarinus rusicola (Melsheimer) 0 0 0 0 0 0 1 1 2 1 2 0 0 0 0 0 7 Labarrus pseudolividus (Balthasar) 0 0 0 0 0 0 0 0 1 1 0 0 2 0 0 0 4 Ataenius strigatus (Say) 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 1 Onthophagus nuchicornis (Linnaeus) 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 1 Totals 325 123 358 400 962 1246 826 556 640 644 618 1042 1022 1377 762 674 11575 147 148 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Fig. 3. Five most abundant dung beetles collected in the Nassawango Creek Preserve from June– October 2009 and March–June 2010. Months have been reorganized to provide general trends. Note the scale is 0–500.

Species previously reported from Canthon chalcites (Haldeman), Geo- Wicomico and/or Worcester County trupes semiopacus Jekel, Onthophagus (Staines 1984), but not collected in orpheus canadensis (Fabricius), and On- our study, include Aphodius ﬁmetarius thophagus orpheus orpheus (Panzer) and (Linnaeus), Ateuchus histeroides Weber, Parataenius simulator Harold (reported Blackburneus stercorosus Melsheimer as Ataenius simulator). We sampled (reported as Aphodius stercorosus), horse ranches on two occasions but did

Fig. 4. Five most abundant dung beetles collected in the Pocomoke State Forest from June–October 2009 and March–June 2010. Months have been reorganized to provide general trends. Note scale is 0–800. VOLUME 114, NUMBER 1 149

Table 4. Species collected in eight sites of Wicomico and Worcester counties from March–December 2010. Speciﬁc bait types are shown to the right (D: human dung, C: carrion, F: fruit, H: horse dung, M: malt, N: no bait). Department of Natural Resources (DNR) specimens were collected from 27 different sites in Worcester County. Specimens marked with an * were also hand collected from different sites. New species not collected in our initial study are indicated with bold.

Species Name Site 1 Site 2 Site 3 Site 4 Site 5 Site 6 Site 7 Site 8 DNR Bait Type

Ateuchus lecontei 1D Blackburneus aegrotus 11 D Canthon pilularius 141 41 15 8 12 24 37 26 66 D C F N Chilothorax distinctus 10 D *Copris minutis 27 25 30 5 28 24 2 9 8 D C F H N Dichotomius carolinus 1D *Geotrupes blackburnii 11 8 5 13 219 11 7 D H Geotrupes egeriei 27 9 6 21 9 13 3 15 D C F M N *Labarrus pseudolividus H Melanocanthon bispinatus 10 17 12 31 33 47 40 D C F M N Onthophagus concinnus 113DN Onthophagus hecate 4 25 11 7 18 13 3 11 D C F M N Onthophagus nuchicornis 1C Onthophagus pennsylvanicus 19 2 5 17 13 6 1 D C F N Onthophagus striatulus 222DN Onthophagus taurus 41 2 D Onthophagus tuberculifrons 67 3348DCFN Phanaeus vindex 124 2 10 12 12 1 1 1 D C F N not have the opportunity to collect from occasionally feces. Many are also found in cattle ranches. This may explain the lack association with bird and mammal nests of these species from our collection ef- (Ratcliffe 1983, Ratcliffe and Paulsen forts. According to Ratcliffe and Paulsen 2008). Though they are commonly referred (2008) and Staines (1984), O. orpheus to as necrophagous beetles, we collected has been collected from cattle dung, bird over 300 specimens of T. hamatus from and mammal nests, and malt traps. Price feces-baited pitfall traps during the ﬁrst (2004, 2006), however, collected this spe- study. Similar ﬁndings were reported in cies in high numbers using cattle dung Price (2004), where over 300 specimens and human feces in an old growth forest were collected in a yearlong study of of central New Jersey. dung beetles attracted to cattle dung in Additional scarabaeoid beetles col- New Jersey. Ratcliffe (1983) made ob- lected during these studies that are not servations of T. hamatus feeding on an reported in Tables 2–4 include one spec- abandoned Canthon brood ball and sug- imen of Ataenius spretulus (Haldeman) gested that this was an opportunistic collected using black light and the fol- occurrence and the event is rare. We lowing Trogidae beetles that were cap- suggest that although most Trogidae do tured in dung and carrion-baited pitfalls: not use feces for food, some species like Omorgus manachus (Herbst), Trox ha- T. hamatus may supplement their diet matus Robinson, Trox terrestris Say, with feces quite frequently. Trox unstriatus Palisot de Beauvois, and European settlement into Maryland Trox variolatus (Melsheimer). Trogidae has changed the landscape from 95% are commonly found feeding on the dry forest cover (Besley 1916) to just 42% in remains of dead animals, feathers, furs, and 2000 (Weber et al. 2006). Fires were set, 150 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON timber destroyed, and land was cleared Manager). This research was in part for agriculture. The original forests were supported by a Salisbury University dominated with hardwoods but are now Guerrieri Undergraduate Summer Re- riddled with pines that have grown up search Endowment. after the land was cleared (Besley 1916). Today, 50% of Wicomico and Worcester Literature Cited counties land cover is used for agriculture Besley, B. S. 1916. The Forests of Maryland. (73% are actually zoned for agriculture) Maryland State Board of Forestry, Forestry (Maryland Department of Natural Re- Pamphlet No. 3. The Advertiser-Republican, sources 2000). Fortunately, there are pro- Baltimore, MD. 152 pp. grams underway that have initiated the Chao, A. 1984. Non-parametric estimation of the number of classes in a population. Scandi- assessment of green infrastructure in navian Journal of Statistics 11: 265–270. Maryland (Weber et al. 2006) and that Estimating the number of shared species in two have begun to convert agricultural land communities. 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User’s Guide and application published at: http://purl.oclc.org/ estimates (accessed May 2011). ACKNOWLEDGMENTS Colwell, R. K. and J. A. Coddington. 1994. Es- We thank Jennifer Frye (Maryland timating terrestrial biodiversity through ex- Department of Natural Resources) for do- trapolation. Philosophical Transactions of the nating specimens from Worcester County. Royal Society (Series B) 345: 101–118. doi:10.1098/rstb.1994.0091 These specimens enhanced our Worcester Gardner, T. A., J. Barlow, I. S. Araujo, T. C. County data. We also thank Jennifer Avila-Pires, A. B. Bonaldo, J. E. Costa, M. C. Frye and Bill Grogan (Salisbury Uni- Esposito, L. V. Ferreira, J. Hawes, M. I. M. versity Emeritus Faculty) for their help Hernandez, M. S. Hoogmoed, R. N. Leite, N. with site determination and suitability. F. Lo-Man-Hung, J. R. Malcolm, M. B. Martins, L. A. M. Mestre, R. Miranda-Santos, We thank Paul Skelley (Florida State W. L. Overal, L. Parry, S. L. Peters, M. A. 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