Surveys of Arkansas dung beetles (Coleoptera: Scarabaeidae and Geotrupidae): phenologies, mass occurrences, state and distributional records.
Abstract: We present the results of an 18 mo survey of the seasonal activity and species composition of dung beetles (Scarabaeidae: Scarabaeinae and Aphodiinae and Geotrupidae: Geotrupes) from a cattle pasture on Crowley's Ridge, Arkansas, and of 17 additional surveys, at various localities and habitats across Arkansas. Collections at the cattle pasture comprised 236,880 beetles, representing 22 species. Labarrus pseudolividus Balthasar comprised 97.7% of the total trap catch, with the majority being collected in Jun., Jul. and Aug. 2007 which involved two separate mass occurrences during that time. From all surveys, we documented 22 state records, of which Colobopterus erraticus (Linnaeus) and Onthophagus taurus (Schreber) represented subtle range extensions to their known distributions, while the remaining 20 species were projected to occur in Arkansas. A checklist, distributional record and bibliography of the 64 species of dung beetles that are now known to occur in Arkansas are presented.
Article Type: Report
Subject: Zoogeography (Research)
Dung beetles (Research)
Dung beetles (Distribution)
Authors: Fiene, Justin G.
Connior, Matthew B.
Androw, Robert
Baldwin, Brian
McKay, Tanja
Pub Date: 04/01/2011
Publication: Name: The American Midland Naturalist Publisher: University of Notre Dame, Department of Biological Sciences Audience: Academic Format: Magazine/Journal Subject: Biological sciences; Earth sciences Copyright: COPYRIGHT 2011 University of Notre Dame, Department of Biological Sciences ISSN: 0003-0031
Issue: Date: April, 2011 Source Volume: 165 Source Issue: 2
Topic: Event Code: 310 Science & research; 690 Goods & services distribution Advertising Code: 59 Channels of Distribution Computer Subject: Company distribution practices
Geographic: Geographic Scope: Arkansas Geographic Name: Arkansas Geographic Code: 1U7AR Arkansas
Accession Number: 254311555
Full Text: INTRODUCTION

By fragmenting and burying excrement, dung beetles (Scarabaeidae: Scarabaeinae and Aphodiinae and Geotrupidae) are an important ecological component in both natural and artificial ecosystems (Bornemissza, 1960; Fincher, 1981; Nichols et al., 2008). Therefore, there is general interest among ecologists and cattle producers in research that aims to conserve and manage these beneficial beetles. Towards this end, to survey dung beetles and catalog their phenologies within a geographic region are the first necessary efforts. Surveys of Scarabaeoidea, which include dung beetles, have been conducted in a number of states (see Kriska and Young, 2002 for review; Riley and Wolfe, 2003). For example, in Texas, the diversity of dung beetles collected in each of seven regions ranged from 44-91 species (Riley and Wolfe, 2003). Limited niche specific surveying has been done in Arkansas (e.g., Ozark pocket gopher, Geomys bursarius ozarkensis Elrod, Zimmerman, Sudman & Heidt, burrows, Kovarik et al., 2008). For the six physiographic regions in Arkansas, only 43 species of dung beetles have been documented to date. Thus, there seemed to be potential to increase the known dung beetle fauna of Arkansas through further surveying efforts of both natural (e.g., forests) and artificial (e.g., cattle pastures) habitats.

We surveyed dung beetles at various localities and habitats across Arkansas to document the dung beetle species and their distributions within the state. The primary study presents the results of an 18 mo survey of dung beetles on Crowley's Ridge. This study is important because it was the first effort to identify dung beetles on Crowley's Ridge in Arkansas, while also being the first survey to document the phenologies and species composition of an assemblage of pasture-inhabiting dung beetles in all of Arkansas.

MATERIALS AND METHODS

Primary locality description.--The primary study was conducted at the Arkansas State University (ASU) Farm Complex inJonesboro, Craighead County, Arkansas. The loessal soil type at the ASU Farm Complex is classified as Loring-Grenada-Henry and is characterized as deep, well- to poorly-drained and slowly permeable (Crow, 1974). The ASU Farm Complex is located on Crowley's Ridge, which is an area of rolling hills that rises abruptly 60-90 m from the flat delta land surrounding it. The Ridge ranges from 1.6 to 16 km wide and runs from southern Missouri to Helena, Arkansas (240 kin-long), encompassing portions of Clay, Greene, Craighead, Poinsett, Cross, St. Francis and Lee Counties. Crowley's Ridge is an erosional remnant of a higher plain from which the present alluvial plain was carved during the Pleistocene interglacial periods when the Mississippi River meandered to the west of the ridge and the Ohio River to the east (Crow, 1974). The vegetation that occurs on Crowley's Ridge is more closely related to the flora (e.g., tulip tree-oak forests) of Tennessee than to the oak hickory forests of the Ozarks, indicating that the biota of Crowley's Ridge had a more recent connection with the Tennessee biota than with the biota of the Ozark/ Ouachita area in Arkansas (Robison and Allen, 1995). Additionally, since Crowley's Ridge has been traditionally used for livestock production and is surrounded by delta that is typically used for crop production (B. Humphrey, pers. comm.), it is a unique area to study dung beetle diversity.

The study site was comprised of two adjacent pastures that were 3.2 and 4.0 ha of open pasture, respectively, each with 0.81 ha of mature hardwood timber. A herd of Angus crossbred cattle, Bos taurus Linnaeus, were rotated between the two pastures, which consisted of tall fescue, Festuca arundinacea Schreber, and Bermuda grass, Cynodon dactylon (Linnaeus). Climatological data were acquired through the Jonesboro Airport weather station that was located 1 km from the study site.

We monitored the seasonal activity of adult dung beetles from 27 Mar. 2007 to 28 Sept. 2008. We evenly spaced ten baited pit-fall traps (-23 m apart) in the open pasture along the fence (253 m) that separated the two pastures. Traps 1 and 10 were ~23 m from the edge of the pasture and the wooded timber, respectively. Traps consisted of two buckets (19.0 cmw X 23.5 cm h), one placed inside the other, with a wire mesh (2.5 cm X 2.5 cm) over the mouth of the trap to support the dung baits. Baits were made of cattle dung (~75 g) wrapped in cheesecloth. The cattle dung was collected fresh (<3 h) and frozen until use (-20 C). We buried the traps in the ground with the lip of the trap level with the soil surface. Then, we added a solution of water (~1 liter), laundry detergent (~50 ml), and table salt (~60 g) to each trap. Vegetation within 0.5 m of the traps was trimmed to less than 5 cm to enhance dispersal of bait odors. Beetles were removed from traps and baits were replaced on a weekly basis throughout most of the study except during winter months (Nov. through early Feb.) when beetles were collected bi-weekly. In total, we collected beetles from 69 trapping periods throughout the 18 mo study. On five occasions a trap was either filled with mud or tipped over and was removed from analysis. Beetles were stored in 70% ethanol until they were sorted, counted and identified. When traps consisted of >6000 beetles, bulk samples of beetles were oven-dried for 5 d at 55 C and then weighed. The number of beetles in the bulk sample was then estimated using the average number of beetles per gram calculated from five subsamples of 0.2 g or 0.5 g.

Phenologies of beetles were examined by pooling the total number of individuals collected from all traps during a baiting period. We separated months in which beetles had been collected from both years to compare annual patterns of species composition.

Additional localities.--To supplement the aforementioned study with an updated state list, dung beeries were also collected at 17 additional localities in Arkansas. Our efforts to collect beetles occurred from 1995-2009 and varied among each locality in the frequency and method of collection. In general, beetles were collected from flight intercept traps, ultraviolet and black lights traps and/or at various types of patchy resources (fruit, carrion, excrement, mushrooms) that were used as baits for pitfall traps or from which beetles were collected in situ. The collection labels are reported in the following format: Month (year), collection method, County (specific locality description), number of specimens collected; additional locality.

Taxonomic identification.--We identified beetles to species level by using taxonomic keys (Robinson, 1948a; Howden, 1955; Howden and Cartwright, 1963; Cartwright, 1974; Edmonds, 1994; Genier, 2000; Ratcliffe et al., 2002; Gordon and Skelley, 2007) with reference to specimens in the Arkansas State University collection, Texas A&M University collection, and the Carnegie Museum of Natural History and personal communication with Dr. Paul Skelley (Florida Department of Agriculture and Consumer Services--Division of Plant Industry) and Ed Riley (Texas A&M University, Department of Entomology).

RESULTS

Primary locality.--We collected a total of 236,880 beetles representing 22 species from the cattle pasture survey on Crowley's Ridge (Table 1). Labarrus pseudolividus Balthasar represented 97.7% of the composition. The majority of L. pseudolividus (211,485 individuals) were collected in the months of Jun., Jul. and Aug. in 2007 (Fig. 1A). The presence of L. pseudolividus occurred from Apr. through Nov. with two periods of high abundance. Each peak lasted for roughly 1 mo, with the first peak starting in mid-Jun, in both years, while the second peak was variable and started in late Jul. in 2007 and late Aug. in 2008.

The seasonal activity for all species at the ASU Farm Complex is reported for those represented by >2 individuals (Figs. 1-5), and those represented by <2 individuals (Table 1). In general, the majority of beetles were collected during the summer when the ambient temperature was greatest and precipitation was lowest (Fig. 6). The average daily maximum temperature for each year (Apr.-Sept.) was 29.9 C in 2007 and 27.3 C in 2008. Total precipitation for each year (Apr.-Sept.) was 33.5 cm in 2007 and 48.3 cm in 2008.

Additional localities.--A total of 30 species of dung beetles were collected from 17 additional localities across Arkansas and are presented as follows:

Aphodius fimetarius (Linnaeus): Jun. (2009) in situ cow dung, Marion Co. (woodland pasture), 4 specimens.

Ataenius apicalis Hinton:Jun. (2002) Ultraviolet (UV) light trap, Pulaski Co. (18.5 km W of Little Rock), 2 specimens; May (2005) UV light, Ouachita Co. (Poison Springs Wildlife Mgt. Area, 8.04 km SE Bluff City), 1 specimen.

Ataenius gracilis (Melsheimer): Jun. (2002) UV light trap, Pulaski Co. (18.5 km W of Little Rock), 4 specimens; Jun. (2005) UV light, Garland Co. (Camp Clear Fork), 3 specimens.

Ataenius hesperius Cartwright: Jun. (2002) UV light trap, Pulaski Co. (18.5 km W of Little Rock), 2 specimens; May (1995) unknown collection method, Pope Co. (Arkansas River at Dardanelle), 6 specimens.

Ataenius imbricatus (Melsheimer): Jun. (2002) UV light trap, Pulaski Co. (18.5 km W of Little Rock), 2 specimens; May (2005) UV light, Ouachita Co. (Poison Springs Wildlife Mgt. Area, 8.04 km SE Bluff City), 6 specimens; Mar. (2009) dung baited pitfall trap in Baird's pocket gopher, Geomys breviceps Baird, burrows via methods described in Skelley and Gordon (2001), Calhoun Co. (cattle pasture, 2 km W Harrel), 2 specimens.

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Ataenius picinus Harold: May (2005) UV light, Ouachita Co. (Poison Springs Wildlife Mgt. Area, 8.04 km SE Bluff City), 2 specimens.

Ataenius platensis (Blanchard): Jul. (2002) UV light trap, Pulaski Co. (18.5 km W of Little Rock), 1 specimen; May (2005) UV light, Ouachita Co. (Poison Springs Wildlife Mgt. Area, 8.04 km SE Bluff City), 1 specimen; May (2005) UV light trap, Montgomery Co. (Ouachita National Forest, 4.5 km NE Norman, Crystal Campground), 1 specimen; Mar. (2009) dung baited pitfall trap in Baird's pocket gopher, C, eomys breviceps Baird, burrows via methods described in Skelley and Gordon (2001), Calhoun Co. (cattle pasture, 2 km W Harrel), 2 specimens.

Ataenius strigatus (Say): Jun. (2002) UV light trap, Pulaski Co. (18.5 km W of Little Rock), 1 specimen;Jul. (2002) UV light trap, Pulaski Co. (18.5 km W of Little Rock), 1 specimen; Jun. (2005) UV light trap, Logan Co. (Mt. Magazine State Park), 1 specimen;Jun. (2005) UV light, Garland Co. (Camp Clear Fork), 1 specimen.

Ateuchus histeroides Weber: Oct. (1998) "intercept trap," Pulaski Co. (Little Rock), 1 specimen;Jun. (2008), UV light trap, Garland Co. (Camp Clear Fork), 1 specimen; 9 May-8 Jun. (2008) four pitfall traps each baited with a woodland vole, Microtus pinetorum LeConte, with each trap separated by -50 m, Craighead Co. (Crowley's Ridge), 31 specimens; Jun. (2009) in situ horse dung, Equus cabaUus L., Searcy Co. (woodland trail), 7 specimens; Jun. (2009) in situ cow dung, Marion Co. (woodland pasture), 3 specimens.

[FIGURE 6 OMITTED]

Blackburneus lentus (Horn): Jun. (2002) UV light trap, Pulaski Co. (18.5 km W of Little Rock), 1 specimen; May (2005) UV light, Ouachita Co. (Poison Springs Wildlife Mgt. Area, 8.04 km SE Bluff City), 1 specimen.

Blackburneus rubeolus (Palisot de Beauvois) : May (2000) UV light trap, Pulaski Co. (Little Rock), 1 specimen.

Blackburneus stercorosus (Melsheimer):Jun. (2002) UV light trap, Pulaski Co. (18.5 km W of Little Rock), 1 specimen; Jun. (2005) UV light, Garland Co. (Camp Clear Fork), 4 specimens.

Canthon chalcites (Haldeman): May (1995) unknown collection method, Searcy Co. (Rt. 14 at Buffalo River), 1 specimen; Jun. (2009) in situ dog dung, Canis lupusfamilians Linnaeus, Marion Co. (rural lawn), 1 specimen.

Copris fricator (Fabricius): Apr.-May (2008) UV light trap, Polk Co. (Mena), 3 specimens.

Cryptoscatomaseter acuminatus (Cartwright): Nov. (2009) excavation of Baird's pocket gopher burrow, Ouachita Co. (road right-of-way), 2 specimens.

Deltochilum gibbosum gibbosum (Fabricius) :Jun.-Oct (1996-1998) flight intercept or carrion traps, Pulaski Co. (Little Rock), 8 total specimens.

Geotrupes blackburnii excrementi Say: Sept.-Jun. (1996-1999) flight intercept or carrion traps, Pulaski Co. (Little rock), 24 specimens; Mar. (1996) unknown collection method, Pulaski Co. (Little Rock), 1 specimen; Apr. (1998) flight intercept trap, Pulaski Co. (Pinnacle Mountain State Park), 1 specimen; Oct. (2008), Polk Co. ("on ground"), 1 specimen.

Geotrupes hornii Blanchard: Jul. (1998) flight intercept trap, Pulaski Co. (Little Rock), 1 specimen; May (2001) flight intercept trap, Pulaski Co. (Little Rock), 1 specimen.

Geotrupes splendidus miarhophagus Say: Sept.-Jul. (1995-1998) flight intercept or carrion traps, Pulaski Co. (Little Rock), 36 total specimens; Jan., May and Oct. (1996) unknown collection method, Pulaski Co. (Little Rock), 4 specimens; Oct. (1996) in situ "dung," Jun. Pulaski Co. (Little Rock), 1 specimen; Sept. (2006), Polk Co. ("garage floor"), 1 specimen; Jun. (2009) in situ horse dung, Searcy (woodland trail), 1 specimen; Jul. (2009) in situ bananas, Polk Co., 1 specimen.

Labarrus pseudolividus Balthasar:Jul. (2002) UV light trap, Pulaski Co. (18.5 km W of Little Rock), 1 specimen; Jun. (2005) UV light trap, Logan Co. (Mt. Magazine State Park), 1 specimen.

Martineziana dutertrei (Chalumeau) : May (2005) UV light trap, Montgomery Co. (Ouachita National Forest, 4.5 km NE Norman, Crystal Campground), 3 specimens; Jun. (2005) uv light trap, Garland Co. (Camp Clear Fork), 1 specimen.

Onthophagus gazella (Fabricius): Aug.-Sept. (1996-1998) black light, black light trap, Pulaski Co. (Little Rock), 6 specimens; May (2005) UV light, Ouachita Co. (Poison Springs Wildlife Mgt. Area, 8.04 km SE Bluff City), 2 specimens.

Onthophagus hecate hecate (Panzer): Mar.-Jul. (1996-1998) malaise trap, flight intercept trap, in situ pig dung, Pulaski Co. (Little Rock), 32 total specimens; 9 May-8 Jun. (2008) four pitfall traps each baited with a woodland vole with each trap separated by -50 m, Craighead Co. (Crowley's Ridge), 1 specimen; Jun. (2009) in situ horse dung, Searcy Co. (woodland trail), 5 specimens; Jun. (2009) in situ cow dung, Marion Co. (woodland pasture), 3 specimens.

Onthophagus medorensis Brown: Apr. (2009) excavation of Baird's pocket gopher burrow, Cleburne Co. (hay pasture), 1 specimen.

Onthophagus pennsylvanicus Harold: Jun. (2009) in situ horse dung, Searcy Co. (woodland trail), 3 specimens.

Onthophagus striatulus striatulus (Palisot de Beauvois): Jun. (1996) in situ mushrooms, Garland Co. (Lake Ouachita St. Pk.), 3 specimens; Jun. (1996) black light trap, Pulaski Co. (Little Rock), 1 specimen; 9 May-8 Jun. (2008) four pitfall traps each baited with a woodland vole with each trap separated by ~50 m, Craighead Co. (Crowley's Ridge), 136 specimens.

Onthophagus subaeneus (Palisot de Beauvois): May (1998) flight intercept, Pulaski Co. (Little Rock), 1 specimen; May (1998) UV light trap, Scott Co. (3.2 km NE Cedar Creek), 1 specimen.

Onthophagus taurus (Schreber): Jun. (2009) in situ horse dung, Searcy Co. (woodland trail), 39 specimens; Jun. (2009) in situ cow dung, Marion Co. (woodland pasture), 3 specimens.

Oscarinus rusicola (Melsheimer): Jun. (2002) UV light trap, Pulaski Co. (18.5 km W of Little Rock), 1 specimen.

Phanaeus vindex MacLeay: May (1995) unknown collection method, Searcy Co. (Rt. 14 at Buffalo River), 1 specimen.

Discussion

The beetle comprising the majority of the samples from the cattle pasture survey, Labarrus pseudolividus, is a small (3.5-5.8 mm), 'dweller' beetle that oviposits and develops 'freely' in the resource (Cambefort and Hanski, 1991). Possibly native to North America, this species is widely distributed (southern United States, south to Chile and Argentina), and commonly collected in southern North America (Gordon and Skelley, 2007). Gordon and Skelley (2007) found that most nearctic specimens previously identified as Labarrus lividus Olivier sensu lato Aphodius lividus, were L. pseudolividus. Therefore, citations of L. lividus will be included; however, in all cases it is not known if the type specimens in these studies agree with the general misidentification of this species.

Using developmental data of Labarrus lividus, which completes development from egg to adult in 25-45 d, the bimodal activity of L. pseudolividus evidenced by the large collection in Jun. and Aug. (Fig. 1), was interpreted as the emergence of the first filial ([F.sub.1]), over-wintering generation in Jun. and the emergence of the second filial generation ([F.sub.2]) in Aug. (Lindquist, 1935). Large collections of dung beetles using baited pitfall traps are not entirely uncommon (Fincher et al., 1986; Hanski and Cambefort, 1991; Floate and Gill, 1998; Bertone et al., 2005). Yet the abundance and dominance of L. pseudolividus was somewhat unique with respect to similar surveys. It seems these abundances were reflective of an event called a mass occurrence that, seemingly, happened in both generations in that year. Such events have been reported for beetle species whose densities reach in the hundreds or thousands of individuals per dung pat (Table 2), while our results described one of these events at a substantially larger scale (7.2 ha study site). Seamans (1934) reported Aphodius distinctus Muller (now Chilothorax distinctus) to mass occur annually in the "countless thousands" in the spring and autumn. The results of this study, conducted over 2 y, indicated greater between-year variation than the descriptions presented by Seamans (1934) and support the observation by Finn and Gittings (2003) that these events occur at a low frequency. Interestingly, mass occurrences have involved species whose larvae exhibit some degree of either obligate or facultative saprophagy (Hanski, 1991; Finn and Gittings, 2003). This holds true for the larval feeding habits of L. pseudolividus (Gordon and Skelley, 2007), and is potentially an important clue for understanding the large abundance collected in 2007 (Hanski, 1991; Finn and Gittings, 2003). For this to be the case, the influx in the developmental resource would have come about in the fall of 2006 (when the adult [F.sub.2] generation of 2006 were ovipositing) and also in Jun. of 2007 when the [F.sub.1] generation were ovipositing. At this point, such a scenario seemingly raises more questions than answers, and highlights the need for additional research. To address the developmental habitat hypothesis for mass occurences and to better understand adult abundances in general, future research should sample for larvae in a variety of developmental habitats i.e., dung pats, soil (K. D. Floate, pers. comm.), near the feeding rings where hay accumulates and leaf litter.

Onthophagus gazella (Fabricius) is an exotic species of dung beetle that was released in northwest Arkansas at two sites in the late 1970s, but failed to establish (Lancaster and Hunter, 1978). In 1983, the species was reported from southwest Arkansas and was thought to have dispersed from Louisiana and/or Texas (Hunter and Fincher, 1985). Assuming that this species is colonizing new localities in a northern direction, it is likely the range extension of O. gazella to northeastern Arkansas did not occur recently since this species was collected in southeastern Missouri in 1997 (MacRae and Penn, 2001). Onthophagus gazella completes development in ~30 d (Blume and Aga, 1978), and could possibly complete 5-6 generations each year being active from Jun. through Nov. in our study. In both years a single peak in collections of O. gazella (encompassing multiple collection dates) was observed (Fig. 2A) suggesting a univoltine natural history; however, this has not been experimentally verified.

We collected two exotic species, Onthophagus taurus and Colobopterus erraticus, that are new state records and represented subtle range extensions from their known distributions. Onthophagus taurus is an exotic species that was accidentally introduced to Florida in 1971 (Fincher and Woodruff, 1975). Since its initial detection, O. taurus had been released in northwest Arkansas where its establishment is uncertain (Lancaster and Hunter, 1978). It has been collected in three neighboring states: central Mississippi, eastern Tennessee and southern Missouri (Lago, 1979; Hoebeke and Beucke, 1997; MacRae and Penn, 2001). Aside from releases in California and Texas, O. taurus appears to be restricted to the eastern United States (Hoebeke and Beucke, 1997). Because O. taurus has not been reported from Iowa, Illinois or Indiana, the populations in southern Missouri possibly originated from Arkansas or Tennessee as a northward expansion. Therefore, three scenarios could have lead to the population of O. taurus on Crowleys Ridge: (1) the Franklin County population in northwestern Arkansas established in 1978 and spread east, (2) the Mississippi/ Tennessee populations expanded north and west, (3) both Franklin County and Mississippi/Tennessee populations merged at an unknown location within the last 29 y, suggesting the Crowley's Ridge population is a mix of both of these populations. In Jun. 2009, O. taurus was collected from the Ozark plateau in southern Marion County and northern Searcy County, Arkansas. This location is approximately half way between Franklin County and Crowley's Ridge. Trapping for O. taurus at the release site in Franklin County would address these hypotheses, especially if no specimens are collected. At our study site, we observed 5-6 peaks of O. taurus (Fig. 1B) that we roughly correlated with 5-6 completed generations given its eight month seasonal activity and developmental time of 4-5 weeks, depending on temperature (Wardhaugh et al., 2001).

Colobopterus erraticus is a species native to Europe that was accidentally introduced to the United States (Gordon and Skelley, 2007). The distribution of this species is Nova Scotia, south to Georgia, west and north to Missouri, Montana and British Columbia (Hunter et al., 1987; Gordon and Skelley, 2007). States between Georgia and Arkansas (Alabama, Mississippi and Tennessee) have no records of C. erraticus and tentatively suggest the Crowley's Ridge population originated in Missouri. Collections of this species occurred from Apr. through mid Jun. (Fig. 3A), and with a developmental time of 6-7 wk (Rojewski, 1983), it is likely C. erraticus is univoltine.

The present study adds 22 dung beetle species to the known insect fauna of Arkansas, 20 of which were projected to occur but not documented (Table 3). This study increased the known dung beetle fauna in Arkansas by 66%, highlighting the importance of additional statewide dung beetle survey, and the investigation of museum collections to produce new records and/or validate suspected occurrences. We also documented a significant collection of Labarrus pseudolividus over a period of about 3 mo using baited pitfall traps, which apparently involved two separate mass occurrences during that time. These events caused the dung pats appearance to be generally flattened, with the periphery having a saw-dust consistency and likely had positive effects on the degradation. Understanding the nature of mass occurrences so that we are able to more actively predict and control these events has convincing potential for increasing the efficiency of cattle and pasture management. Future research on the dung beetles in Arkansas has potential for elucidating a poorly understand phenomenon called a mass occurrence and for the documentation of additional species and their distributions within the state.

Acknowledgments.--We thank Paul Skelley (Florida Department of Agriculture and Consumer Services--Division of Plant Industry) and Ed Riley (Texas A&M University) for confirmation of species identifications, Kevin Floate (Agriculture and Agri-Food Canada, Alberta, Canada) and two anonymous reviewers for comments on an earlier version of this manuscript and Mike Johnson (Arkansas State University) for technical support at the research site. This project was partially funded by the Arkansas Biosciences Institute and the Arkansas Game and Fish Commission. Voucher specimens were placed in the Arkansas State University Museum, Carnegie Museum of Natural History and the Florida State Museum Collection of Arthropods.

SUBMITTED 11 JANUARY 2010

ACCEPTED 15 SEPTEMBER 2010

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WOODRUFF, R. E. 1973. The scarab beetles of Florida (Coleoptera: Scarabaeidae). Part I. The Laparosticti (Subfamilies Scarabainae, Aphodiinae, Hybosorinae, Ochodaeinae, Geotrupinae, Acanthocerinae). Arthropods of Florida and Neighboring Land Areas, Vol. 8:1-222.

JUSTIN G. FIENE (1) AND MATTHEW B. CONNIOR (2)

Department of Biological Sciences, Arkansas State University, State University 72467

ROBERT ANDROW

Carnegie Museum of Natural History, Section of Invertebrate Zoology, 4400 Forbes Avenue, Pittsburgh,

Pennsylvania 15213

BRIAN BALDWIN

439Fence Road, Jessieville, Arkansas 71949

AND

TANJA McKAY (3)

Department of Biological Sciences, Arkansas State University, State University 72467 and Department of

Entomology, University of Arkansas, Fayetteville 72701

(1) Present address: Department of Entomology, Texas A&M University, College Station 77845

(2) Present address: South Arkansas Community College, El Dorado 71730

(3) Corresponding author: e-mail: tmckay@astate.edu
TABLE 1.--Number of dung beetles (Scarabaeidae: Scarabaeinae
and Aphodiinae and Geotrupidae: Geotrupes) trapped and species
composition (% of total catch) in 2007 and 2008, (Apr.-Sept.)
respectively, and for the entire 18 mo survey (27 Mar. 2007-28
Sept. 2008) from primary survey at cattle pasture in Jonesboro,
Arkansas

                                             Number of beetles
                                               (% of total)

                                                Apr.-Sept.

Species                                            2007

Aphodius fimetarius (Linnaeus) (a)                3 (0.01)
Ataenius platensis Blanchard                    191 (.09)
At. spretulus (Haldeman)                        274 (.13)
Blackburneus stercorosus (Melsheimer)             1 (<0.01) (c)
Calamosternus gmnarius (Linnaeus) (a)             8 (<0.01)
Colobopterus erraticus (Linnaeus) (b)            51 (.02)
Dichotomius carolinus (Linnaeus)                    ...
Geotrupes blackburnii excrementi Say              1 (<0.01)
Labarrus pseudolividus Balthasar            214,030 (98.57)
Lechorodius lutulentus (Haldeman)                   ...
Le. terrninalis (Say)                             1 (<0.01)
Melinopterus femoralis (Say)                     13 (<0.01)
Onthophagus gazella (Fabricius) (b)              60 (.03)
O. hecate hecate (Panzer)                      1355 (.62)
O. pennsylvanicus Harold                        570 (.26)
O. taurus (Schreber) (a)                        522 (.24)
O.scarinus rusicola (Melsheimer)                  1 (<0.01) (c)
Otophorus haemomhoidalis (Linnaeus) (a)           5 (<0.01)
Pseudagolius bicolor (Say)                          ...
Phanaeus vindex MacLeay                           8 (<0.01)
Platytomus longulus (Cartwright)                 37 (.02)
Pseudocanthon perpLxus (LeConte)                  1 (<0.01) (g)
TOTAL                                       217,132

                                             Number of beetles
                                               (% of total)

                                                Apr.-Sept.

Species                                            2008

Aphodius fimetarius (Linnaeus) (a)                3 (0.01)
Ataenius platensis Blanchard                     53 (.27)
At. spretulus (Haldeman)                        102 (.53)
Blackburneus stercorosus (Melsheimer)               ...
Calamosternus gmnarius (Linnaeus) (a)            25 (.13)
Colobopterus erraticus (Linnaeus) (b)           257 (1.33)
Dichotomius carolinus (Linnaeus)                  1 (<0.01) (d)
Geotrupes blackburnii excrementi Say                ...
Labarrus pseudolividus Balthasar             17,471 (90.51)
Lechorodius lutulentus (Haldeman)                   ...
Le. terrninalis (Say)                               ...
Melinopterus femoralis (Say)                      1 (<0.01)
Onthophagus gazella (Fabricius) (b)             369 (1.91)
O. hecate hecate (Panzer)                       437 (2.26)
O. pennsylvanicus Harold                        237 (l.23)
O. taurus (Schreber) (a)                        325 (l.68)
O.scarinus rusicola (Melsheimer)                  1 (<0.01) (f)
Otophorus haemomhoidalis (Linnaeus) (a)           7 (0.04)
Pseudagolius bicolor (Say)                          ...
Phanaeus vindex MacLeay                           8 (0.04)
Platytomus longulus (Cartwright)                  4 (0.02)
Pseudocanthon perpLxus (LeConte)                    ...
TOTAL                                        19,301

                                             Number of beetles
                                               (% of total)

Species                                        Entire study

Aphodius fimetarius (Linnaeus) (a)               11 (<0.01)
Ataenius platensis Blanchard                    244 (0.1)
At. spretulus (Haldeman)                        376 (0.16)
Blackburneus stercorosus (Melsheimer)             1 (<0.01)
Calamosternus gmnarius (Linnaeus) (a)            61 (0.02)
Colobopterus erraticus (Linnaeus) (b)           308 (0.13)
Dichotomius carolinus (Linnaeus)                  1 (<0.01)
Geotrupes blackburnii excrementi Say             27 (0.01)
Labarrus pseudolividus Balthasar            231,626 (97.74)
Lechorodius lutulentus (Haldeman)                 1 (<0.01) (e)
Le. terrninalis (Say)                             7 (<0.01)
Melinopterus femoralis (Say)                     15 (<0.01)
Onthophagus gazella (Fabricius) (b)             431 (0.18)
O. hecate hecate (Panzer)                      1890 (0.8)
O. pennsylvanicus Harold                        813 (0.34)
O. taurus (Schreber) (a)                        973 (0.41)
O.scarinus rusicola (Melsheimer)                  2 (<0.01)
Otophorus haemomhoidalis (Linnaeus) (a)          12 (<0.01)
Pseudagolius bicolor (Say)                       14 (<0.01)
Phanaeus vindex MacLeay                          16 (<0.01)
Platytomus longulus (Cartwright)                 50 (0.02)
Pseudocanthon perpLxus (LeConte)                  1 (<0.01)
TOTAL                                       236,880

(a) Accidentally introduced species (Fincher and Woodruff, 1975;
Gordon and Skelley, 2007)

(b) Intentionally introduced species (Blume and Aga, 1978)

Collection dates: (c) (5-5-07); (d) (8-3-08); (e) (11-17-07);
(f) (5-25-08); (g) (7-31-07)

Beetles collected from Oct. 2007-Mar. 2008 are represented as
the difference in the total number of beetles collected in the
entire study compared to beetles collected from Apr.-Sept. of
both years

TABLE 2.--Mass occurrences of dung beetles reported by species,
number of individuals per pat and location

                                            Individuals
Reference                 Species             per pat      Location

Lindquist (1935)   Labarrus lividus            1113       Texas
                   (Olivier)

Mohr (1943)        Chilothorax distinctus      1097       Illinois
                   Muller

Merritt (1974)     Aphodius fimetarius        200-600     California
                   (Linnaeus)

Holter (1982)      Nimbus contaminates         3273       Denmark
                   (Herbst)

TABLE 3.--A checklist, distributional record and bibliography of dung
beetles (Scarabaeinae: Scarabaeinae and Aphodiinae and Geotrupidae:
Geotrupes) in Arkansas

Species                           County                 Citation

Aphodius fimetarius       Craighead, Marion         present study
(Linnaeus) (a)

Ataeniopsis figurator     Drew, Crittenden,         Cartwright (1974)
(Harold)                  Hempstead, Jackson,
                          Lafayette

A. rugopygus              Hempstead                 Cartwright (1974)
(Cartwright)

Ataenius apicalis         Columbia, Jackson,        Cartwright (1974);
Hinton                    Ouachita, Pulaski,        present study
                          Stuttgart, Washington

A. gracilis               Benton, Garland,          Cartwright (1974);
(Melsheimer)              Pulaski, Washington       present study

A. imbricatus             Benton, Calhoun,          Cartwright (1974);
(Melsheimer)              Hempstead, Ouachita,      present study
                          Pulaski, Sebastian,
                          Washington

A. hesperius              Pope, Pulaski             present study
Cartwright (b)

A. oklahomensis Brown     Washington, Craighead     Cartwright (1974)

A. ovatulus Horn          State citation only       Cartwright (1974)

A. picinus Harold         Columbia, Drew,           Cartwright (1974)
                          Hempstead, Lafayette,
                          Ouachita

A. platensis              Calhoun, Craighead,       Cartwright (1974);
(Blanchard)               Crawford, Johnson,        present study
                          Logan, Montgomery,
                          Ouachita, Pike,
                          Pulaski, Washington

A. robustus Horn          Hempstead, Lonoke,        Cartwright (1974)
                          Pulaski, Washington

A. spretulus              Arkansas, Columbia,       Cartwright (1974);
(Blanchard)               Craighead, Drew,          present study
                          Garland, Hempstead,
                          Madison, Pulaski, Pike,
                          Randolph, Washington

A. texanus Harold         Crittenden, Hope,         Cartwright (1974)
                          Jackson County,
                          Washington

A. wenzelii Horn          Hope                      Cartwright (1974)

Ateuchus histeroides      Marion, Searcy,           present study
Weber (c)                 Craighead, Garland,
                          Pulaski

Blackburneus lentus       Ouachita, Pulaski         present study
(Horn) (a)

Blackburneus rubeolus     Pulaski                   present study
(Palisot de Beauvois)
(a)

Blackbumeus stercorosus   Craighead, Garland,       present study
(Melsheimer) (a)          Pulaski

Calamosternus granarius   Craighead                 present study
(Linnaeus) (a)

Canthon chalcites         Marion, Searcy            present study
(Haldeman) (e)

C. depressipennis         State citation only       Robinson (1948a)
LeConte

C. pilularius             Hempstead, Miller,        Halffter (1961)
(Linnaeus)                Mississippi,
                          Montgomery, Pulaski,
                          Sharp

Colobopterus erraticus    Craighead                 present study
(Linnaeus) (f)

Copris fricator           Arkansas, Benton,         Matthews (1961);
(Fabricius)               Lawrence, Logan, Polk     present study

C. minutus (Drury)        Garland, Hempstead,       Matthews (1961)
                          Pike

Cryptoscatomaseter        Ovachita Co.              present study
acuminatus

C. haldemani (Horn)       Izard                     Kovarik et al.
                                                    (2008)

C. oklahomensis (Brown)   Izard                     Gordon and Skelley
                                                    (2007); Kovarik
                                                    et al. (2008)

Dellacasiellus kirni      Izard                     Kovarik et al.
(Cartwright)                                        (2008)

Deltochilum gibbosum      Pulaski                   present study
gibbosum (Fabricius)
(b)

Dichotomius carolinus     Craighead                 present study
(Linnaeus) (e)

Geomyphilus insolitus     Izard                     Gordon and Skelley
(Brown)                                             (2007); Kovarik
                                                    et al. (2008)

Geotrupes blackbumii      Craighead, Franklin,      present study;
excrementi Say            Hempstead, Lawrence,      Howden (1955)
                          Washington, Pulaski

G. homii Blanchard        Bentonville, Pulaski      Howden (1964);
                                                    present study

G. opacus Haldeman        Hempstead, Lawrence,      Howden (1955)
                          Marion

G. spLndidus              Mississippi, Polk,        Howden (1955);
miarhophagus Say          Pulaski, Searcy           present study

Irrasinus stupidus        "Ark., south west"        Gordon and
(Horn)                                              Skelley (2007)

Labarrus pseudobvidus     Craighead, Logan,         present study
Balthasar (a)             Pulaski

Lechoradius lutuLntus     Craighead                 present study
(Haldeman) (a)

L. terminalis (Say) (a)   Craighead                 present study

Martineziana dutertrei    Montgomery, Garland       present study
(Chalumeau) (b)

Melanocanthon             State citation only       Robinson (1948a)
nigricornis (Say)

Melinopterus femoralis    State citation only,      Gordon and
(Say)                     Craighead                 Skelley (2007);
                                                    present study

Onthophagus               Washington                Howden and
cavemicollis Howden                                 Cartwright (1963)
and Cartwright

O. gazella (Fabricius)    Craighead, Lafayette,     present study;
                          Miller, Ouahcita,         Hunter and
                          Pulaski                   Fincher (1985)

O. hecate hecate          SW, SE, Ncentral AR,      Howden and
(Panzer)                  Craighead, Lonoke,        Cartwright (1963);
                          Pulaski                   present study

O. medorensis Brown       Boone                     Howden and
                                                    Cartwright (1963)

O. oklahomensis Brown     Miller, Lawrence          Howden and
                                                    Cartwright (1963)

O. orpheus pseudorpheus   Lonoke                    Howden and
Howden and Cartwright                               Catrwright (1963)

O. pennsylvanicus         NW, SW and Central AR,    Howden and
Harold                    Craighead                 Cartwright (1963);
                                                    present study

O. striatulus             NW and SW AR,             Howden and
striatulus (Palisot de    Craighead, Garland,       Cartwright (1963);
Beauvois)                 Pulaski                   present study

O. subaeneus (Palisot     Pulaski, Scott            present study
de Beauvois) (b)

O. taurus (Schreber)      Craighead, Marion,        present study
(f)                       Searcy

O. tuberculifrons         Washington                Howden and
Harold                                              Cartwright (1963)

Oscarinus rusicola        Craighead, Pulaski        present study
(Melsheimer) (a)

Otophorus                 Craighead                 present study
haemorrhoidalis
(Linnaeus) (a)

Phanaeus difformis        Franklin, Jefferson       Edmonds (1994)
LeConte

P. triangularis           Carroll, Little River,    Robinson (1948b)
triangularis (Say)        Madison, Washington

P. vindex MacLeay         Craighead,                present study;
                          Independence,             Edmonds (1994)
                          Jefferson, Johnson,
                          Lafayette, Lawrence,
                          Polk, Searcy,
                          Washington

Pseudagolius bicolor      Craighead                 present study
(Say) (a)

Pseudocanthon perplexus   State citation only,      Blanchard (1885);
(LeConte)                 Craighead                 present study

Platytomus longulus       Craighead                 present study
(Caxwright) (b)

Scabrostomus sepultus     Izard                     Gordon and Skelley
(Cartwright)                                        (2007); Kovarik
                                                    et al. (2008)

(a) Verified state record, projected to occur by Gordon and
Skelley (2007)

(b) Expected occurrence based on published distribution

(c) Verified state record, projected to occur by Genier (2000)

(d) Verified state record, projected to occur by Cartwright (1974)

(e) Verified state record, projected to occur by Woodruff (1973)

(f) State records not expected to occur in Arkansas
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