Open access journal: http://periodicos.uefs.br/ojs/index.php/sociobiology ISSN: 0361-6525 DOI: 10.13102/sociobiology.v63i1.764Sociobiology 63(1): 623-627 (March, 2016) Geographic range of Pachycondyla harpax (Fabricius) (Hymenoptera: Formicidae) Introduction Ponerine ants (subfamily Ponerinae) include >1000 species (Bolton 2015), many of which are well known for their potent sting. For example, Brachyponera sennaarensis (Mayr) (formerly Pachycondyla sennaarensis) is a widespread species of sub-Saharan Africa and the Middle East that is well known for its powerful sting, which sometimes leads to anaphylactic shock in humans and even death (Wetterer, 2013). Brachyponera chinensis (Emery) (formerly Pachycondyla chinensis) is an East Asian species now spreading through the eastern US that has a painful sting that can also induce severe allergic reaction (Guénard & Dunn, 2010). Here, I examine the geographic range of the ponerine ant Pachycondyla harpax (Fabricius). Wittenborn and Jeschke (2011) included P. harpax on a list of exotic ant species established in North America. Based on Wittenborn and Jeschke (2011), Miravete et al. (2013; now retracted) listed P. harpax as Neotropical species from South Abstract Pachycondyla harpax (Fabricius) is a widespread and conspicuous New World ponerine ant (subfamily Ponerinae). To examine the geographic distribution of P. harpax, I compiled and mapped published and unpublished specimen records from >1600 sites. I documented the earliest known P. harpax records for 28 geographic areas (countries, West Indian islands, and US states), including four for which I found no previously published records: the islands of Guadeloupe, Margarita, and Tobago and the US state of Georgia. Pachycondyla harpax has been recorded from every country in South and Central America except Chile and Uruguay. Pachycondyla harpax is also now known from six West Indian islands: Grenada, Guadeloupe, Jamaica, Margarita, Trinidad, and Tobago. The known continental range of P. harpax appears to be essentially continuous, extending from Rio Grande do Sul, Brazil in the south (31.8°S) to Wood County, Texas in the north (32.8°N), including the continental islands of Margarita, Tobago, and Trinidad. Isolated island populations of P. harpax on Grenada, Guadeloupe, and Jamaica may be exotic, introduced through human commerce. In the US, it is unclear why P. harpax populations are only known from Texas, Louisiana, and Georgia, when there would appear to be suitable habitat for this species all along the Gulf coast of Alabama, Mississippi, and Florida. Sociobiology An international journal on social insects JK Wetterer Article History Edited by Flavia Esteves, CAS, USA Evandro Nascimento Silva, UEFS, Brazil Received 05 February 2015 Initial acceptance 07 April 2015 Final acceptance 04 September 2015 Publication date 29 April 2016 Keywords Biogeography, native range, native species, Ponerinae, West Indies. Corresponding author James K. Wetterer Wilkes Honors College Florida Atlantic University, 5353 Parkside Drive, Jupiter, FL, 33458, USA E-Mail: wetterer@fau.edu America that is an introduced and established exotic species in the US. Pachycondyla harpax is a large (up to 10 mm in length), conspicuous New World ponerine ant (Fig 1). Pachycondyla harpax has an exceptionally broad geographic range in South, Central, and North America (Wheeler, 1900) and occupies a great diversity of natural and disturbed environments. Mackay and Mackay (2010) wrote that P. harpax is “found in a wide variety of habitats, ranging from dry forest, late dry season gap forest, urban environments, parks, grassy areas, coffee, cacao and banana plantations, cypress swamps, to oak forests, oak riparian forests, rocky wet quebradas, arid scrub, palm thorn forests, tropical deciduous forests, tropical evergreen forests, second growth tropical forests, transitional bamboo/ cloud forest, old growth dry tropical forest, steep rocky forest slopes, ridge forest, wet mountain forests, riparian rain forest, montane evergreen forest, lowland forests, cloud forest and riparian tropical rain forests.” Longino (2001) wrote: “I have RESEARCH ARTICLE - ANTS Wilkes Honors College, Florida Atlantic University, Jupiter, Florida, USA JK Wetterer – Geographic range of Pachycondyla harpax (Fabricius)624 never found a nest of this common species. The nest must be subterranean. If they nested in the leaf litter or in dead wood, nests would be more frequently encountered.” Taxonomy and identification is absent on the dorsum of the meso soma and the petiole is rectangular shaped with a distinct dorsal face and the posterior lateral margin forms a sharp carina, which is barely evident as it passes to the anterior edge of the petiole.” Brown (1950) suggested that P. harpax may actually be a species complex rather than a single species. Schmidt (2013) determined that the genus Pachycondyla, as formerly constituted, was polyphyletic. In Schmidt and Shattuck’s (2014) revision, however, P. harpax remained in the genus Pachycondyla along with ten other species. Methods Using published and unpublished records, I documented the worldwide range of P. harpax. I obtained unpublished site records from museum specimens in the collections of the Museum of Comparative Zoology (MCZ) and the Smithsonian Institution (SI). In addition, I used on- line databases with collection information on specimens by Antweb (www.antweb.org), and the Global Biodiversity Information Facility (www.gbif.org). I received unpublished collection information of P. harpax records from C. Alatorre- Bracamontes (Mexico) and E. Mendoza (El Salvador). Finally, I collected P. harpax specimens from Costa Rica, El Salvador, Guadeloupe, Margarita, Trinidad, and Tobago. I obtained geo-coordinates for collection sites from published references, specimen 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 I had no other record for this region, I used the coordinates of the capital or largest town within the region or, in the case of small islands and natural areas, the center of the region. Published records usually included collection dates, but when this was not the case, I was sometimes 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. For example, Nils Holmgren visited in Peru and Bolivia in 1904-1905, thus bracketing the date he collected P. harpax specimens examined by Wheeler (1925). Henry T. Vanderford (1901-1990), who collected the only known P. harpax specimen from Georgia (identification confirmed by S.P. Cover), worked for the Georgia Department of Agriculture until at least 1967 (Williams, 1986). Results I compiled and mapped published and unpublished specimen records from >1600 sites. I documented the earliest known P. harpax records for 28 geographic areas (countries, West Indian islands, and US states), including four for which I found no previously published records: the islands of Guadeloupe, Margarita, and Tobago and the state of Georgia. Pachycondyla harpax has been recorded from every Fig 1. Pachycondyla harpax. a) head and b) lateral view of worker from Costa Rica (CASENT0249149; P.S. Ward leg.; photos by R. Perry from antweb.org). Fabricius (1804) described Formica harpax (= P. harpax) from South America. Junior synonym include Pachycondyla harpax concinna Wheeler (from Bolivia, Brazil, and Peru), Pachycondyla harpax dibullana Forel (from Colombia), Pachycondyla harpax irina Wheeler (from Colombia, Costa Rica, Guatemala, Guyana, and Peru), Pachycondyla montezumia Smith (from Mexico), Pomera amplinoda Buckley (from Texas), and Pachycondyla orizabana Norton (from Mexico) (see Brown, 1950). Mackay and Mackay (2010) wrote that the P. harpax: “worker can be separated from most other species in the genus, as it lacks the malar carina, the eye is small but is located less than one maximum diameter from the anterior edge of the head. The pronotal carina is poorly developed but forms a shiny raised line (usually), the metanotal suture Sociobiology 63(1): 623-627 (March, 2016) 625 country in South and Central America except Chile and Uruguay (Table 1). Pachycondyla harpax is also now known from six West Indian islands: Grenada, Guadeloupe, Jamaica, Margarita, Trinidad, and Tobago (Table 2). Most records of P. harpax come from intact forest habitats. In 2015, antweb.org recorded that P. harpax was “found most commonly in these habitats: 382 times found in mature wet forest, 169 times found in tropical moist forest, 152 times found in tropical rainforest, 120 times found in tropical wet forest, 106 times found in secondary lowland rainforest, 106 times found in lowland wet forest, 104 times found in montane wet forest, 77 times found in secondary wet forest, 51 times found in lowland rainforest, 50 times found in mesophil forest.” I have collected P. harpax at 42 sites, 39 of which came from inland forest sites on West Indian islands: Guadeloupe (4), Margarita (1), Tobago (10), and Trinidad (24). The three others came from Central America: Costa Rica (beach vegetation in Tortuguero National Park) and El Salvador (two gardens in urban San Salvador). Discussion Wheeler (1900) wrote that P. harpax ranged from “Texas through Mexico, Central America and Brazil to Bolivia and Paraguay.” More recent records extend that range to include Louisiana and Georgia in the north, Argentina in the south, plus several West Indian islands (Table 1). The native continental range of P. harpax appears to be essentially continuous from Rio Grande do Sul, Brazil (31.8°S) in the south to Wood County, Texas (32.8°N) in the north (Fig 2). Although there are numerous apparent gaps in this continental range (e.g., in central Brazil), it is likely that much of this is an artifact of insufficient sampling. It is unclear why P. harpax populations in the US are only known from Texas, Louisiana, and Georgia when there would appear to be suitable habitat for this species below 31°N all along the Gulf coast of Alabama and Mississippi and into Florida. Populations of P. harpax on the West Indian islands of Margarita, Trinidad, and Tobago represent part of the continuous continental range because these are continental islands that were connected to South America when sea levels were >100 m lower 15,000-30,000 years ago (Lambeck et al., 2014). The West Indian islands of Grenada, Guadeloupe, and Jamaica, however, have never been connected to the adjacent continent. It is unclear how P. harpax population came to colonize Grenada, Guadeloupe, and Jamaica. It is possible that the populations of P. harpax on these islands are exotic, introduced through human commerce. On all three islands, however, P. harpax was collected almost exclusively in intact native forest, an atypical locale for an exotic species. Genetic analyses could be useful in determining whether P. harpax is native or exotic on Grenada, Guadeloupe, and Jamaica. I found no evidence indicating that P. harpax is exotic to continental North America. In addition to P. harpax, Wittenborn and Jeschke (2011) appear to have misclassified numerous other ant species as exotics to North America that are actually natives. For example, Gnamptogenys hartmani (Wheeler), Labidus coecus (Latreille), and Leptogenys elongata (Buckley), all have distributions in the southern US that appear South America ≤1804 (Fabricius, 1804) Mexico ≤1858 (Smith, 1858 as P. montezumia) Texas ≤1866 (Buckley, 1866 as Pomera amplinoda) Colombia ≤1870 (Mayr, 1870) Guatemala ≤1883 (Forel, 1899) Venezuela 1887-1888 (Emery, 1890a) Costa Rica 1889 (Emery, 1890b) Bolivia 1885-1893 (Emery, 1894) Brazil ≤1895 (Forel, 1895) Guyana ≤1895 (Forel, 1895) Paraguay ≤1895 (Forel, 1895) Nicaragua ≤1899 (Forel, 1899) Peru 1904-1905 (Wheeler, 1925 as P. harpax irina & P. harpax concinna) Belize 1905-1906 (Wheeler, 1907) Panama 1911 (W.M. Wheeler, MCZ): Ancón Louisiana 1913 (J.R. Horton, SI): Nairn Honduras 1920 (Mann, 1922) Ecuador 1922 (F.X. Williams, SI): Baños El Salvador 1959 (N. Krauss, SI): Santa Ana Surinam 1959 (Kempf, 1961) Argentina ≤1978 (Kusnezov, 1978) +Georgia ≤1990 (H.T. Vanderford, SI): Eastman French Guiana 1996 (G.D. Alpert & M. Moffett, MCZ): Paracou Forest Table 1. Earliest known records for Pachycondyla harpax in South, Central, and North America. + = no previously published records. MCZ = Museum of Comparative Zoology. SI = Smithsonian Institution. Jamaica 1909 (Wheeler, 1911) Grenada 1912-1913 (R. Thaxter, MCZ): Grand Etang Trinidad 1913 (Wheeler, 1916) +Tobago 2003 (J.K. Wetterer, MCZ): Gilpin Trace +Guadeloupe 2008 (J.K. Wetterer, MCZ): Carbet Falls Road +Margarita 2010 (J.K. Wetterer, MCZ): SE of San Sebastian Table 2. Earliest known records for Pachycondyla harpax on West Indian islands. JK Wetterer – Geographic range of Pachycondyla harpax (Fabricius)626 to be the northern end of continuous native ranges and give no indication of being exotic to North America. Other species that Wittenborn and Jeschke (2011) most likely misclassified as exotics include Cephalotes varians (Smith), a widespread arboreal species in Cuba, the Bahamas, and Florida (Andrade & Baroni Urbani, 1999) and Leptogenys manni (Wheeler), a species endemic to Florida (Trager & Johnson, 1988). Mark Deyrup (pers. comm.), who has encyclopedic knowledge of the ants of Florida, confirmed that he knows of no reason why these two species would be that considered exotic in Florida (the only US state where they occur). Regrettably, Wittemborn and Jeschke (2011) did not provide evidence to support their classification. There is some danger that if native US species, such as P. harpax, are erroneously considered to be invasive exotic species, they may be treated as such and exterminated, rather than valued and protected. Acknowledgments I thank M. Wetterer for comments on this manuscript; S. Cover and J. Longino for ant identification; S. Cover (MCZ), T. Schultz (SI), and M. Deyrup (ABS) for help with their respective ant collections; C. Alatorre-Bracamontes and E. Mendoza for unpublished records; 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; Florida Atlantic University and the National Science Foundation (DES-0515648) for financial support. References Andrade, M. L. de and Baroni Urbani, C. (1999). Diversity and adaptation in the ant genus Cephalotes, past and present. Stuttgarter Beiträge zur Naturkunde - Serie B, 271: 1-889. Bolton, B. (2015). An online catalog of the ants of the world. http://www.antcat.org/catalog/430052. (accessed date: 4 September, 2015). Brown, W.L., Jr. (1950). Morphological, taxonomic, and other notes on ants. The Wasmann Journal of Biology, 8: 241-250. Buckley, S.B. (1866). Descriptions of new species of North American Formicidae. Proceedings of the Entomological Society of Philadelfia, 6: 152-172. Emery, C. (1890a). Voyage de M. E. Simon au Venezuela (Décembre 1887 - Avril 1888). Formicides. Bulletin de la Societe Entomologique de France (6) 10: 55-76. Emery, C. (1890b). Studi sulle formiche della fauna neotropica. Bollettino della Societa Entomologica Italiana, 22: 38-80. Emery, C. (1894). Studi sulle formiche della fauna neotropica. Bollettino della Societa Entomologica Italiana, 26: 137-241. Fabricius, J.C. (1804). Systema Piezatorum secundum ordines, genera, species, adjectis synonymis, locis, observationibus, descriptionibus. Brunswick: C. Reichard, 439 p Forel, A. (1895). A fauna das formigas do Brazil. Boletim do Museu Paraense de Historia Natural e Ethnographia, 1: 89-139. Forel, A. (1899). Biologia Centrali-Americana; or, contributions to the knowledge of the fauna and flora of Mexico and Central America. Insecta. Hymenoptera. 3. Formicidae. London: R.H. Porter, 160 p Guénard, B. & Dunn, R.R. (2010). A new (old), invasive ant in the hardwood forests of eastern North America and its potentially widespread impacts. PLoS ONE 5 (7): e11614. DOI:10.1371/journal.pone.0011614 Kempf, W.W. (1961). A survey of the ants of the soil fauna in Surinam (Hymenoptera: Formicidae). Studia Entomologica, 4: 481-524. Kusnezov, N. and Golbach, R. (1978). Hormigas argentinas: clave para su identificación. Ministerio de Cultura y Educacion, Fundación Miguel Lillo, Tucuman, Miscelanea, 61, 147 p. Lambeck, K., Roubya, H., Purcella, A., Sunc, Y. & Sambridgea, M. (2014). Sea level and global ice volumes from the Last Glacial Maximum to the Holocene. PNAS, 111: 15296-15303. Longino, J.T. (2001). Pachycondyla harpax (Fabricius). http:// academic.evergreen.edu/projects/ants/genera/Pachycondyla/ SPECIES/harpax/harpax.html Mackay, W.P. & Mackay, E.E. (2010). The systematics and biology of the New World ants of the genus Pachycondyla (Hymenoptera: Formicidae). Lewiston: Edwin Mellon Press, 642 p. Fig 2. Geographic distribution of Pachycondyla harpax records. Sociobiology 63(1): 623-627 (March, 2016) 627 Mann, W.M. (1922). Ants from Honduras and Guatemala. Proceedings of the United States National Museum, 61: 1-54. Mayr, G. (1870). Formicidae novogranadensis. Sitzungsberichte der Kaiserlichen Akademie der Wissenschaften. Mathematisch- naturwissenschaftliche Classe. Abteilung I 61: 370-4177. Miravete, V., Roura-Pascual, N., Dunn, R.R. & Gómez, C. (2013). How many and which ant species are being accidentally moved around the world? Biology Letters, 9: 20130540 (in data supplement). doi: 10.1098/rsbl.2013.0540 Schmidt, C.A. (2013). Molecular phylogenetics of ponerine ants (Hymenoptera: Formicidae: Ponerinae). Zootaxa, 3647: 201-250. doi: 10.11646/zootaxa.3647.2.1 Schmidt, C.A. and Shattuck, S.O. (2014). The higher classification of the ant subfamily Ponerinae (Hymenoptera: Formicidae), with a review of ponerine ecology and behavior. Zootaxa, 3817: 1-242. doi: 10.11646/zootaxa.3817.1.1 Smith, F. (1858). Catalogue of hymenopterous insects in the collection of the British Museum. Part VI. Formicidae. London: British Museum, 216 p Trager, J.C. and Johnson, C. (1988). The ant genus Leptogenys (Hymenoptera: Formicidae, Ponerinae) in the United States. In Trager, J.C. (Ed.), Advances in Myrmecology (pp. 29-34). Leiden: E. J. Brill. Wetterer, J.K. (2013). Geographic spread of the samsum or sword ant, Pachycondyla (Brachyponera) sennaarensis (Hymenoptera: Formicidae). Myrmecological News, 18: 13-18. Wheeler, W.M. (1900). A study of some Texas Ponerinae. Biological Bulletin, 2: 1-31. Wheeler, W.M. (1907). A collection of ants from British Honduras. Bulletin of the American Museum of Natural History, 23: 271-277. Wheeler, W.M. (1911). Additions to the ant fauna of Jamaica. Bulletin of the American Museum of Natural History, 30: 21-29. Wheeler, W.M. (1916). Ants collected in British Guiana by the expedition of the American Museum of Natural History during 1911. Bulletin of the American Museum of Natural History, 35: 1-14. Wheeler, W.M. (1925). Neotropical ants in the collections of the Royal Museum of Stockholm, Part 1. Arkiv för Zoologi, 17: 1-55. Williams, D.F. (1986). Chemical baits: specificity and effects on other ant species. In: C.S. Lofgren and R.K. Vander Meer (Eds.), Fire ants and leaf-cutting ants: biology and management (pp 378-386). Boulder: Westview Press. Wittenborn, D. & Jeschke, J.M. (2011). Characteristics of exotic ants in North America. NeoBiota 10: 47-64 (Appendix 1). doi: 10.3897/neobiota.10.1047.app1