Notes on the Scarabaeoid Dung Beetles (Coleoptera: Scarabaeidae, Geotrupidae, and Trogidae) of Hutcheson Memorial Forest, New Jersey, U.S.A.1

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Volume 117, Number 3, May and June 2006 347

SCIENTIFIC NOTE

NOTES ON THE SCARABAEOID DUNG BEETLES (COLEOPTERA: SCARABAEIDAE, GEOTRUPIDAE, AND TROGIDAE) OF HUTCHESON MEMORIAL FOREST, NEW JERSEY, U.S.A.1

Dana L. Price2

Hutcheson Memorial Forest (HMF), Somerset Co., New Jersey, is one of the last uncut forests in the Mid-Atlantic States and is apparently the only uncut upland forest in New Jersey (Hutcheson Memorial Forest, 2004). The old forest consists of a 26-ha mixed-oak stand believed to be relatively undisturbed since 1702 (Buell et al., 1954; Buell, 1957), and is surrounded by more than 70-ha of young forest, abandoned agricultural fields undergoing plant succession, and a number of adjoining 1-ha abandoned fields (Robertson and Vitousek, 1981; Hutcheson Memorial Forest, 2004). The two primary objectives of the HMF cen- ter, since it was established in 1955 are to: 1) maintain the old woods in their natural state, and 2) study the natural ecological and evolutionary processes present. Consequently, the tract has become one of the most intensively studied ecological areas in the world (Buell and Forman, 1982). This paper is part of an ongoing study of the ecology of dung beetles at this site. In 2002 – 2003 the seasonal abundance and species diversity of scarabaeoid dung beetles were examined in the old forest and a disturbed field in HMF (Price, 2004). A total of 24 species were captured. Here I present a follow-up study using dung and carrion as bait in order to assess variation in food preference. The col- lection of new species, introduced species, and food preference (coprophagous vs. necrophagous) are discussed. Dung beetles were collected at two HMF sites: the old-growth forest (N40° 29.753' W74°33.852') and the disturbed field (N40°30.028' W74°33.835'). Pitfall trap collections were made using two types of dung (dog feces and human feces) and two kinds of carrion (chicken legs and fish) as bait (see Price, 2004 for trap details). These baits were used due to their availability and freshness. Two arrays of four pitfalls each were placed in each site (total of eight pitfalls per site). The four traps of each array were each set at a corner of a 10 m square, and the two arrays at each site were set 20 m apart. Collections were made three times during the summer of 2004 (July 3 to July 5; July 31 to August 2; and August 31 to September 2), each for a 48-hour period. A total of 1023 dung beetles, representing 15 species were collected (Table 1), two of which have not previously been reported in HMF: Aphodius stercorosus, ______1 Received on August 28, 2005. Accepted on October 28, 2005. 2 Graduate Program of Ecology and Evolution, Rutgers University, DEENR, First Floor, 14 College Farm Road, New Brunswick, New Jersey 08901 U.S.A. E-mail: [email protected]. Mailed on July ??, 2006 348 ENTOMOLOGICAL NEWS from both the forest and field sites, and Bolboceras liebecki from the field site. This genus has not previously been reported from HMF. To date, a total of 28 species have been reported from HMF (Shubeck et al., 1977; Price, 2004; the present study). Price (2004) reported Geotrupes semiopacus and Onthophagus striatulus to be captured only in the old forest. In the present study, one specimen of each species was collected in the disturbed field as well. Additional beetle families captured are shown in Table 2. Histeridae, Silphi- dae, and Staphylinidae were by far the most abundant families. According to Downie and Arnett (1996) histerid beetles in carrion and dung beetle communi- ties are predaceous on other insects and their larvae. Food preference for species collected were determined with SAS (1990), using Proc GLM. Log transformation was used to correct for non-homogeneous variances. Onthophagus hecate, a species common in both the forest and field sites was attracted to all four types of bait (Table 1), though it had a significant preference for dung over carrion (P < 0.0002; F = 8.15). Species only collected in dung-baited traps that showed a significant preference for human feces over dog feces include; Onthophagus pennsylvanicus (P < 0.0002; F = 10.42), O. orpheus (P < 0.0027; F = 6.65), Phanaeus vindex (P < 0.0162; F = 4.36), and Dialytes truncatus (reported as Aphodius sp. in Price, 2004: P < 0.0022; F = 6.95). Onthophagus striatulus appeared to show a preference for carrion over dung (7/7 individuals were collected in carrion traps), though it was not statisti- cally significant (P < 0.1176; F = 2.07). In Price’s (2004) study, O. striatulus was captured in pitfalls baited with cow dung, but they were not given a choice of carrion verses dung. Trox hamatus, associated with chicken feathers, squirrel carrion and fox bur- rows (Vaurie, 1955), was also collected in large numbers (299 specimens) from pitfalls baited with cow dung (Price, 2004). Because Trox species in general are thought to be mainly carrion feeding (dried animal parts), and are only occa- sionally collected on dung, I suggest that individuals collected from the pitfalls with cow dung may have been attracted to the mucky water in the pitfalls (the water and bait were replaced only once a week). However, in this study, only 3 specimens were attracted to the carrion, 15 to dog feces and 2 to human feces. Further sampling might support the observation that this species has a preference for dung over carrion. Price (2004) reported two introduced species collected in HMF: Onthophagus taurus and Aphodius distinctus. Both species were collected only in the disturbed field site. During this study, Onthophagus taurus was captured in both the old- growth forest (1 specimen) and disturbed field site (2 specimens). Aphodius distinctus was not collected at either location. Volume 117, Number 3, May and June 2006 349

Table 1. A list of all scarabaeoid dung beetles (Coleoptera: Scarabaeidae, Geotrupidae, and Trogidae) collected from carrion (chicken and fish), and dung (dog and human) during the summer of 2004.

Field Forest Total Species C F D H C F D H

Aphodius bicolor Say 2 1 3 Aphodius rubripennis Horn 3 3 Aphodius stercorosus Melsheimer 1 1 2 Bolbocerus liebecki (Wallis) 1 1 Copris minutis (Drury) 6 10 20 26 5 3 9 79 Dialytes truncates (Melsheimer) 29 29 Geotrupes semiopacus Jekel 1 1 8 26 36 Onthophagus hecate (Panzer) 8 20 118 113 1 6 84 74 424 Onthophagus orpheus (Panzer) 10 75 85 Onthophagus pennsylvanicus Harold 106 209 315 Onthophagus striatulus (Palisot de Beauvois) 1 1 5 7 Onthophagus taurus (Shreber) 2 1 3 Phanaeus vindex Macleay 2 14 16 Trox hamatus Robinson 2 1 10 2 3 18 Trox sp. 1 1 2

Total 18 31 256 369 2 17 111 219 1023 350 ENTOMOLOGICAL NEWS

Table 2. Additional families collected from dung and carrion-baited pitfall traps during the summer of 2004. Field Forest Total Family C F D H C F D H Carabidae 3 3 1 28 30 10 1 76 Chrysomedlidae 1 1 Curculionidae 1 1 Elateridae 1 1 2 Histeridae 11 139 22 28 4 4 4 212 Hydrophilidae 1 1 *Scarabaeidae 2 1 3 Silphidae 72 176 97 15 28 30 22 12 452 Staphylinidae 29 140+ 57 70 56 131 86 50 479 Total 112 319 182 116 112 197 122 67 1227 * Three specimens of Rutelinae (not dung beetles).

ACKNOWLEDGEMENTS I am grateful to my adviser, Michael May and to W.D. Edmonds for reviewing this manuscript. I thank Francois Genier (Canadian Museum of Nature, Ottawa, Ontario, Canada) for his identification and confir- mation of Aphodius species in this manuscript. I would also like to thank Colleen Hatfield and my labmate Jessica Ware for discussion and valuable input. This research was supported by the William L. Hutcheson Memorial Forest Summer Research Grant.

LITERATURE CITED Buell, M. F. 1957. The mature oak forest of Mettler’s Woods. William Hutcheson L. Memorial Forest Bulletin 1: 16–19. Buell, M. F., H. F. Buell, and J. A. Small. 1954. Fire in the history of Mettler’s Woods. Bulletin of the Torrey Botanical Club 81: 253–255. Buell, M. F. and Forman. 1982. Three decades of research at Hutcheson Memorial Forest, New Jersey (USA). The William L. Hutcheson Memorial Forest Bulletin 6(2): 24–32. Downie, N. M. and R. H. Arnett, Jr. 1996. The Beetles of Northeastern North America. Volume I: Introduction; Suborders Archestemata, Adephaga, and Polyphaga, thru Superfamily Cantharoidea. The Sandhill Crane Press, Gainsville Florida. Hutcheson Memorial Forest. 2004. Website hosted by Rutgers University: http//www.rci.edu/~deenr/ HMF.html (accessed November 2004). Price, D. L. 2004. Species diversity and seasonal abundance of scarabaeoid dung beetles (Coleoptera: Scarabaeidae, Geotrupidae, and Trogidae) attracted to cow dung in central New Jersey. Journal of the New York Entomological Society 112(4): 334-347. Robertson, G. P. and P. M. Vitousek. 1981. Nitrification in primary and secondary succession. Ecology 62: 376–386. SAS Institute Inc. 1990. SAS/STAT User’s Guide, Version 6, Fourth Edition, Volume 2. SAS Institute Inc. Cary, North Carolina, U.S.A. Shubeck, P. P., N. M. Downie, R. L. Wenzel, and S. B. Peck. 1977. Species composition of carrion beetles in a Mixed-Oak Forest. William L. Hutcheson Memorial Forest Bulletin 4(1): 12-17. Vaurie, P. 1955. A revision of the genus Trox in North America (Coleoptera, Scarabaeidae). Bulletin of the American Museum of Natural History 106(1): 1-371.