geographic spread of pyramica hexamera (hymenoptera

© Koninklijke Brill NV, Leiden, 2012 DOI 10.1163/187498311X615754

Terrestrial Arthropod Reviews 5 (2012) 3–14 brill.nl/tarT A R

Geographic spread of Pyramica hexamera (Hymenoptera: Formicidae: Dacetini) in the

southeastern USA

Joe A. MacGown1,* and James K. Wetterer2 1Department of Entomology and Plant Pathology, Mississippi State University,

Mississippi State, Mississippi 39762 USA 2Wilkes Honors College, Florida Atlantic University, 5353 Parkside Drive,

Jupiter, Florida 33458 USA *Corresponding author; e-mail: [email protected]

Received: 11 October 2011; accepted 27 October 2011

SummaryPyramica hexamera is a tiny predatory ant that feeds on minute soil arthropods. Originally from East Asia, P. hexamera has been recently introduced to North America, apparently through human commerce. Here we document the known range of P. hexamera in Asia and the New World. We compiled and mapped 73 site records of P. hexamera, all from East Asia and the southeastern US. In Asia, P. hexamera records range from 21.9°N to 36.4°N (earliest date and number of sites in parentheses) in Japan (1949; 25), South Korea (1982; 2), and Taiwan (1992; 6). In the US, P. hexamera records range from 28.6°N to 34.3°N: in Florida (1987; 2), Louisiana (1987; 5), Mississippi (2003; 32), and Alabama (2007; 1). Pyramica hexam-era is reported for the first time from Alabama.© Koninklijke Brill NV, Leiden, 2012

KeywordsDacetine ants; biogeography; biological invasion; exotic species; invasive species

Introduction

Dacetines (Hymenoptera: Formicidae: Dacetini) are predatory ants that generally feed on tiny soil arthropods (Wilson, 1953). Members of this group are small, cryptically colored, and slow moving. They rarely forage openly above ground and become motionless when disturbed. As a result, most people, including field biologists, remain unaware of their presence even in areas where they are common. For example, Carroll (2009) surveyed 15 bayhead tree-islands in the Florida Everglades and did not find any dacetines in visual searches, yet found dacetines in Berlese extractions of leaf litter from 14 of the 15 islands.

4 J.A. MacGown and J.K. Wetterer / Terrestrial Arthropod Reviews 5 (2012) 3–14

Over 900 species and subspecies of dacetine ants are currently recognized (Bolton et al. 2007). Of the 37 dacetines reported from the southeastern US, four are Old World exotics (Deyrup and Cover, 2009; MacGown and Hill, 2010). Three of these have already achieved broad distributions in both the Old World and the New World, spread through human commerce: Strumigenys emmae (Emery), Strumigenys rogeri Emery, and Pyramica membranifera (Emery) (Wetterer, 2011). In the US, only P. mem-branifera is naturalized across the southern states; S. emmae and S. rogeri are naturalized only in peninsular Florida (Wetterer, 2011). In this paper we document that the fourth species, Pyramica hexamera (Brown), is spreading across the southeastern US.

The first reported collection of P. hexamera outside its native Asia was in 1987 from Marion County, Florida (Deyrup, 1988). This species was again collected in Florida in 1998 in Hernando County (Deyrup, 2003), but has not been reported from the state since that time. Pyramica hexamera was also collected in West Feliciana Parish, Louisiana in 1987 (Bolton, 2000). Subsequently, it was collected from another site in West Feliciana Parish in 1999, in St. Tammany Parish in 2000, East Baton Rouge Parish in 2003, and Iberia Parish in 2003 (Dash, 2004). The first reported collections from Mississippi were from Chickasaw (2003, 3 sites), Lauderdale (2003, 1 site), and Jones Counties (2004, 2 sites) (MacGown et al., 2005).

To examine the geographic spread of P. hexamera, we compiled and mapped speci-men records from its native and exotic range.

Taxonomy and identification

Brown (1958) described Epitritus hexamerus (= P. hexamera) from two sites in Japan. Bolton (1999) transferred E. hexamerus to Pyramica. Baroni Urbani and De Andrade (2007) synonymized Pyramica with Strumigenys. More recently, Bolton and Alpert (2011) reconfirmed Pyramica as a valid genus.

Pyramica hexamera is a minute (~2.0 mm in total length), brown ant with elongate mandibles, each of which terminates with a long apicodorsal tooth (Figure 1A,B). Pyramica hexamera has spongiform tissue present ventral to the petiole and postpetiole (Figure 2A), and large, scale-like setae present on much of the dorsal surface of the body except the gaster (Figures 1A,B and 2A-D). Pyramica hexamera is unlikely to be confused with any other related dacetine in the southeastern US (Deyrup and Cover, 2009).

Biology

Pyramica hexamera is a specialized ambush predator of small long-bodied soil arthro-pods, such as Diplura, Chilopoda, and Collembola (Masuko, 1984). Masuko (2009) reported, “Diplura composed 60% of the prey in the field material.” This prey prefer-ence is unusual compared with most dacetines whose biologies have been studied, and which apparently prey primarily on Collembola (Masuko, 2009). Foraging P. hexamera workers hunt for their prey in small crevices in the soil. Upon encountering prey, P. hexamera moves to a crouching position, pulls its antennae back into recessed

J.A. MacGown and J.K. Wetterer / Terrestrial Arthropod Reviews 5 (2012) 3–14 5

antennal scrobes lining the side of the head, closes its mandibles, and remains motion-less. The ant may remain still for >20 min while it waits for the prey to advance and crawl on top of its head. Pyramica hexamera is uniquely equipped for this situation, having a flattened head and slightly upturned mandibles (Figure 1A), each of which

Figure 1. Pyramica hexamera worker: (A) profile view, and (B) full-face view.


terminates with a sharp apical tooth (Figure 2B,C), allowing the ant to strike at prey passing overhead. Additionally, the entire dorsal surface of the head is covered with large scale-like setae, which may be sensory in nature (Figure 2B,D). When the prey is in the correct position, the ant opens its mandibles and suddenly snaps them shut, impaling the prey with the apical teeth (Masuko, 1984). Pyramica hexamera has also been reported to coat its body with soil and other detritus using its forelegs, a behavior that may camouflage the ant’s odor from the prey (Masuko, 1984).

Methods

Using published and unpublished records; we documented the worldwide range of P. hexamera. We obtained unpublished site records from museum specimens in the

Figure 2. Pyramica hexamera worker: (A) spongiform tissue beneath petiole and postpeti-ole, (B) full-face view showing scales covering head, scapes, and mandibles, (C) close-up view of mandibles showing scales and elongate apicodorsal teeth, and (D) close-up of scales on head.


collections of Archbold Biological Station (ABS, identified by M. Deyrup), the Mississippi Entomological Museum (MEM, identified by J. MacGown), the Museum of Comparative Zoology (MCZ, identified by S. Cover), and the Smithsonian Institution (SI, identified by B. Bolton). In addition, we used on-line databases with collection information on specimens by the Essig Museum of Entomology at the University of California, Berkeley (UCB), Antweb (www.antweb.org), and the Global Biodiversity Information Facility (www.gbif.org).

We obtained geo-coordinates for collection sites from published references, speci-men labels, maps, or geography web sites (e.g., earth.google.com, www.tageo.com, and www.fallingrain.com). If a site record listed a geographic region rather than a “point locale,” and we had no other record for this region, we used the coordinates of the larg-est town within the region or, in the case of small islands and natural areas, the center of the region. We did not map records of P. hexamera found in newly imported goods or intercepted in transit by quarantine inspectors. Published records usually included collection dates. In a number of cases, publications did not include the col-lection dates for specimens, but we were able to determine the approximate date based on information on the collector’s travel dates or limit the date by the collector’s date of death.

Results

We compiled and mapped P. hexamera specimen records from a total of 73 sites (Table 1; Figure 3). In Asia, P. hexamera records range from 21.9°N to 36.4°N in Japan, South Korea, and Taiwan. In the US, P. hexamera records range from 28.6°N to 34.3°N: in Florida, Louisiana, Mississippi, and Alabama (Figure 4).

Figure 3. Map showing the worldwide distribution of Pyramica hexamera.


Most records we found of P. hexamera from Asia, where this species of ant is native, are from Japan (25 sites) (Bolton, 2000; Brown, 1958; Masuko, 2009; Ogata and Onoyama, 2003; Sato et al., 2010; Shimazaki and Miyashita, 2005; Terayama and Kihara, 1998; Yamauchi and Ogata, 1995). Only six records have been reported from Taiwan (Anonymous, 1999; Terayama et al., 1995; Terayama and Kubota, 1989) and only two from South Korea (Kim et al., 1992; Lyu et al., 2001; Park and Kim 2002).

We compiled 40 site records of P. hexamera from the US, all from the Southeast. By far, the majority of records of this species from the southeastern US have been from Mississippi (32 site records). Since P. hexamera was first reported from six sites in Mississippi in 2003 (MacGown et al., 2005), collections have been made at 26 addi-tional sites in 17 counties: Adams County: St. Catherine Creek, 31°21'15"N, 91°25'17"W, 11-25 April 2006, B. A. Lotz, Lindgren funnel trap baited with Typosan [Typosan ® is a bark beetle aggregation pheromone for various bark beetles (Coleoptera: Curculionidae: Scolytinae)] (2 alate queens); Natchez State Park, 31°35'49"N, 91°12'03"W, 8 July 2005, J. G. Hill, J. A. MacGown, litter on slope in mixed pine/hardwood forest (1 worker). Chickasaw County: Camp Tik-a-witha, 30°58'15"N, 88°55'12"W, 30 May 2006, R. J. Jones, J. A. MacGown, Berlese-litter at base of Quercus sp. (1 dealate queen). Clarke County: Clarkco State Park, 32°06'05 N, 88°41'59"W, 2 July 2010, J. A. MacGown, J. G. Hill, litter from bases of trees and logs in pine/hardwood forest (2 workers). Clay County: Waddell, 33°34'51"N, 88°52'17"W, 5-25 April 2006, J. F. Standley, Lindgren funnel trap baited with Typosan (1 alate queen). Forrest County: 3.5 mi SSW Dixie, 31°09'57"N, 89°19'19"W, 13 April 2004, B. A. Lotz, Lindgren funnel trap (1 alate female); Brooklyn, 31°03'01"N, 89°11'43"W, G. Christodoulou, 21 June 2005, Berlese-litter in mixed oak-pine forest (1 worker). Franklin County: Homochitto Nat. Forest, 31°30'57", 91°07'41"W, 8 July 2005, J. A. MacGown, J. G. Hill, Berlese-soil under pine log in mixed pine-hardwood forest (2 workers, 1 dealate queen). George County: [no locality data], 2-16 April 2008, D. Haynes, Lindgren funnel trap (3 alate queens). Harrison County: 6 miles NE Wool Market, 30°32'38"N, 88°57'07"W, 23 Mar-13 April 2004, B. A. Lotz, Lindgren funnel trap (1 alate queen). Jasper County: 3 mi E of exit 118

Table 1. Earliest known records for Pyramica hexamera from East Asia and the United States.

# of sites Earliest record

Japan 25 1949 (Brown 1958)Taiwan 6 1982 (Terayama and Kubota 1989)South Korea 2 <1992 (Kim et al. 1992)Florida 2 1987 (Deyrup 1988)Louisiana 5 1987 (Bolton 2000)Mississippi 32 2003 (MacGown et al. 2005)+Alabama 1 2007 (J. Lasiter, MEM)+ = no previously published records.MEM = Mississippi Entomological Museum.


X I59, 31°56'25"N, 88°57'26"W, 21 June 2005, G. Christodoulou, Berlese-hardwood canopy, kudzu groundcover (1 worker). Jefferson County: Natchez Trace Parkway, 31°43'25"N, 91°10'32"W, 8 July 2005, J. A. MacGown, J. G. Hill, Berlese-litter in mixed pine-hardwood forest (3 workers). Jefferson Davis County: Jeff Davis Lake, 31°33'47"N, 89°50'39"W, 11 August 2005, J. A. MacGown, J. G. Hill, Berlese-litter in mixed pine-hardwood forest (1 worker). Lauderdale County: 2 mi WNW Lizelia, 32°32'39"N, 88°41'49"W, 26 April-13 May 2004, J. F. Standley, Lindgren funnel trap (1 alate queen); Meridian, 32°20'39"N, 88°38'47"W, 7 March 2009, J. G. Hill, in mixed forest with undulating topography (3 alate queens); Lauderdale, 32°31'07" N, 88°33'01"W, 6 June 2007, J. G. Hill, L. C. Majure, in bottomland forest (1 dealate queen); Bonita Lakes Reserve, 32°20'33"N, 88°38'41"W, 15 August 2008, J. G. Hill, Berlese-litter in undulating mixed pine-hardwood forest (9 workers, 1 dealate queen); same data except, 32°21'13"N, 88°39'36"W, 25 April 2008, J. G. Hill, J. A. MacGown (2 workers), Berlese-leaf litter in hardwood forest; same data except, J. A. MacGown, J. G. Hill, at base of Carya sp. in hardwood forest (1 dealate queen). Lee County: Natchez Trace Parkway, mi 252, 34°09'01"N, 88°49'04"W, 20 October 2009,

Figure 4. Map showing the known records of Pyramica hexamera in the United States.


J. G. Hill, Berlese-litter from oak-hickory forest in Black Belt Prairie (1 worker). Lowndes County: Lake Lowndes State Park, 33°26'24"N, 88°18'12"W, 9 June 2010, J. G. Hill, J. A. MacGown, litter from base of Quercus sp. in mixed pine-hardwood forest (1 dealate queen). Pontotoc County: Trace State Park, 34°15'24"N, 88°53'12"W, 18 May 2006, J. G. Hill, in mixed forest in Black Belt Prairie (1 dealate queen); Natchez Trace, mi. 247.5, 34°05'49"N, 88°51'38"W, 28 April-5 May 2003, T. Schiefer, J. MacGown, Lindgren funnel trap in deciduous forest (1 alate queen); Natchez Trace Parkway, mi. 247, 34°05'52"N, 88°51'37"W, 20 October 2009, J. G. Hill, Berlese-litter from edge of Black Belt Prairie and oak-hickory forest (1 worker). Scott County: Golden Memorial State Park, 32°34'12"N, 89°24'25"W, 27 April 2006, J. G. Hill, J. A. MacGown, in mixed forest with open areas (15 workers, 1 dealate queen). Wilkinson County: Clark Creek Natural Area, 31°04'17"N, 91°30'41"W, 7 July 2005, J. A. MacGown, J. G. Hill, Berlese-litter in hardwood forest (8 workers).

This species was not included as occurring in Alabama in a recent publication that documented ants occurring in that state (MacGown and Forster, 2005). However, with collections of P. hexamera from Florida, Louisiana, and Mississippi, the likelihood of finding it in Alabama seemed high. Indeed, a single alate queen of this species was collected on 4 April 2007 in a Lindgren funnel trap in Stockton, Baldwin County, Alabama (30°59'N 87°51'W) by J. Lasiter (in MEM collection). This collection repre-sents the first, and only, report of P. hexamera from Alabama. More extensive collecting in the state could result in further records.

Discussion

Ogata and Onoyama (2003) reported that in Japan P. hexamera is a “rare species nest-ing in the soil of broadleaf forests.” Likewise, most collections of P. hexamera from Mississippi were from soil and litter samples in deciduous forests, especially in areas with undulating topography. One reason for its apparent abundance in Mississippi as compared to Florida and Louisiana may be that this species prefers wooded habitats with more varied topography. However, the relatively high number of collections of this species in Mississippi may simply reflect the intensive fieldwork done by the MEM during the past 10 years.

This species may be generally overlooked because unlike most other dacetines, it appears to live in soil rather than leaf litter (Masuko, 1984). Deyrup and Deyrup (1999) extracted litter samples from a wide variety of habitats around Florida and found no P. hexamera specimens in 908 samples. The MEM conducted widespread surveys of ants from 2001 through 2011 in Mississippi, especially in natural habitats on public lands, such as state parks and national forests (MacGown, 2011); whereas, major surveys in Florida were done primarily prior to 2001 (Deyrup et al., 1989; Deyrup 2003). Given that the earliest records of P. hexamera in North America were in 1987, (Florida, Marion County, 3 mi N. Pedro, 19 June 1987, Walter Suter and Louisiana, Iberia Parish, New Iberia, 19 August 1987, W&E Mackay, pitfall trap in forest), it may not have been well established in Florida at the time when major survey work was being done.


Pyramica hexamera appears to have different dietary habits from the three other tramp dacetine ants, S. emmae, S. rogeri, and P. membranifera, all three of which appear to prey primarily on Collembola. Differences in body size and mandible morphology among these three species (and dacetines in general) may be related to differences in the size and type of prey attacked by each species (Hölldobler and Wilson, 1990). The ecological importance of P. hexamera, as well as that of all other dacetine ants, remains largely unstudied.

According to Deyrup et al. (2000), all nine of the exotic dacetine species that have been reported from the southeastern US were probably introduced into the region in soil of nursery stock. This seems especially likely in the case of the soil nesting P. hex-amera, which is native to Japan and Korea. A wide variety of Japanese trees, shrubs, vines, and other plants have been successfully introduced either intentionally or unin-tentionally into the southeastern United States (i.e. Japanese honeysuckle, Lonicera japonica Thunb., Caprifoliaceae; Japanese persimmon, Diospyros kaki Thunb., Ebenaceae; Kudzu, Pueraria lobata Ohwi, Fabaceae; Japanese wisteria, Wisteria flori-bunda D.C., Fabaceae; Japanese climbing fern, Lygodium japonicum Sw., Lygodaceae; Japanese maple, Acer palmatum Thunb., Sapindaceae; and others). Many of these and other Asian plant species now proliferate in the southeastern US, which is climatologi-cally and latitudinally similar to parts of Japan and other Asian regions. Due to its diminutive size and cryptic coloration, queens and/or entire colonies of P. hexamera could have easily have been transported to the US in soil of potted plants undetected.

It is unclear when and where P. hexamera first entered the US. Given that the first records of P. hexamera in the United States were from 1987 in both Louisiana (Bolton, 2000) and Florida (Deyrup, 1988), it is possible that multiple introductions have occurred. For that matter, it could just as likely have been already been well established in Louisiana and/or Mississippi before 1987, but not reported due to lack of collecting during that time period. Prior to 2001, there was a dearth of recent ant records from Louisiana, Mississippi, and Alabama. Moser and Blum (1960) published a preliminary list of ants of Louisiana that did not include records of P. hexamera. However, serious ant faunal studies were not undertaken again until 2001, when Dash began making collections throughout the state as part of his thesis project (Dash, 2004; Dash and Hooper-Bùi, 2007). In addition to the early record from 1987 (Bolton, 2000), Dash reported four new collections from the state (Dash, 2004). Similarly, Marion Smith conducted wide-scale faunal surveys in Mississippi in the early 1900’s as part of a fed-eral mandate to fight the Argentine ant, Linepithema humile (Mayr), (Smith, 1924 a, b, c; 1927; 1928 a, b; 1931 a, b; 1932). Smith did not collect P. hexamera during his studies, and he only reported a total of ten dacetine species from Mississippi. No serious surveys were made in the state again until 2001, when the MEM began inten-sive surveys throughout the state as part of a USDA-ARS Areawide Management of Imported Fire Ant Project. During this project, 27 dacetine species, including P. hexamera, were reported by MacGown and Hill (2010). Because of the great lapse of time between these projects, we do not know when P. hexamera first arrived in the state. Pyramica hexamera was not reported in an early survey of ants of Alabama (Murphree, 1947), nor was it reported from a recent study of ants of Alabama conducted by MacGown and Forster (2005).


Acknowledgments

We thank M. Wetterer for comments on this manuscript; S. Cover for help, encour-agement, and ant identification; S. Cover (MCZ) and M. Deyrup (ABS) for help with their respective ant collections; W. O’Brien for GIS help; D.P. Wojcik and S.D. Porter for compiling their valuable FORMIS bibliography; R. Pasos and W. Howerton of the FAU library for processing so many interlibrary loans; and JoVonn Hill (MEM), Beth Lotz, and Jay Standley (USDA-APHIS) for their collections of ants. The interesting records of alate P. hexamera in Lindgren funnel traps were made possible by USDA-APHIS and the Cooperative Agricultural Pest Survey. This project was supported by Florida Atlantic University, the National Science Foundation (DES-0515648), Mississippi Agricultural and Forestry Experiment Station State Project MIS-311080, and the USDA-ARS Areawide Management of Imported Fire Ant Project (Richard L. Brown, Principal Investigator). Approved for publication as Journal Article No. J-12085 of the Mississippi Agricultural and Forestry Experiment Station, Mississippi State University.

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geographic spread of pyramica hexamera (hymenoptera

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