The Coexistence
Myrmecological News 12 97-108 Vienna, September 2009
Worldwide spread of the destroyer ant, Monomorium destructor (Hymenoptera: For- micidae)
James K. WETTERER
Abstract
The destroyer ant, Monomorium destructor (JERDON, 1851), is a pest in many tropical and subtropical areas, where it is notorious for chewing through the insulation of electrical wires, living in and destroying electrical equipment, and at- tacking people. To evaluate the spread of M. destructor, I compiled published and unpublished specimen records from > 600 sites worldwide. I documented the earliest known M. destructor records for 107 geographic areas (countries, is- land groups, major Caribbean islands, and US states), including many locales for which I found no previously published records: Bahamas, Barbados, Barbuda, Bonaire, Comoro Islands, Curaçao, Grenada, Haiti, Honduras, Kenya, Nevis, Paki- stan, Réunion, St. Lucia, St. Martin, St. Vincent, Tobago, and Venezuela. Monomorium destructor most closely resem- bles several African Monomorium species, and has a seemingly continuous distribution from North Africa to Southeast Asia, suggesting M. destructor originated in North Africa, but is also native to the Middle East and South Asia. Mono- morium destructor is most common as a pest in disturbed arid and semi-arid habitats in the tropics and subtropics. Out- breaks of M. destructor often appear to be fairly localized and short-lived. This pattern of population explosion followed by decline should be taken into consideration in any large-scale efforts to control these ants. Key words: Biogeography, biological invasion, exotic species, Formicidae, invasive species. Myrmecol. News 12: 97-108 (online 17 February 2009)xxx 2008) ISSN 1994-4136 (print), ISSN 1997-3500 (online) Received 29 July 2008; revision received 17 December 2008; accepted 17 December 2008
Prof. Dr. James K. Wetterer, Wilkes Honors College, Florida Atlantic University, 5353 Parkside Drive, Jupiter, FL 33458, USA. E-mail: [email protected]
Introduction Numerous ant species have spread around the world through concluded that M. destructor probably originated in Asia human commerce. Among the best known and most de- (EMERY 1893a, WHEELER 1906, MAYOR 1922, KUSNEZOV structive are the African big-headed ant, Pheidole mega- 1949, SMITH 1965, DLUSSKY & ZABELIN 1985, BOLTON cephala (FABRICIUS, 1793) (WETTERER 2007), the little fire 1987, FOWLER & al. 1994, COLLINGWOOD & al. 2004). ant, Wasmannia auropunctata (ROGER, 1863) (WETTERER For example, COLLINGWOOD (1962) reported M. destructor & PORTER 2003), the red imported fire ant, Solenopsis in- (as junior synonym Monomorium gracillimum (SMITH, victa BUREN, 1972 (ALLEN & al. 2004), and the Argentine 1861)) from Kambaiti, Burma, and concluded "the species ant, Linepithema humile (MAYR, 1868) (HUMAN & GOR- would seem to be truly indigenous in Burma since the spe- DON 1997). While surveying ants on tropical Atlantic and cimens were collected in a mountain area." RUZSKII (1907) West Indian islands, I encountered numerous outbreaks of wrote that M. destructor (as M. gracillimum) probably ori- another destructive pest ant, Monomorium destructor (JER- ginated in Southeast India, and from there, spread West to DON, 1851). For example, on the Cape Verde Islands, many Africa through Persia, Syria, and Arabia, as well as south- people readily shared stories about how these ants recently ern Europe and Polynesia. BERNARD (1944, 1948) wrote appeared in their homes, living in their walls and appli- that M. destructor (as M. gracillimum) was originally from ances, and eating virtually anything around the house, in- the steppes and deserts of central Asia and was in the pro- cluding food, insulation on electrical wires, and the fabric cess of invading the entire Sahara and surrounding regions. of furniture and clothing. Numerous Cape Verde residents BERNARD (1958, 1960) gave a more precise origin, writ- reported being attacked by these ants in their gardens and ing that M. destructor (as M. gracillimum) was introduced homes, and even while in bed. In areas of outbreak in Cape from Iran to North Africa in the 19th century. Verde, M. destructor was well known to locals as the "ninja In contrast, FOREL (1909) reported M. destructor (as ant." Here, I evaluate the worldwide spread and possible M. gracillimum) among native plants in a desert area near geographic origin of M. destructor. Biskra, Tunisia, and speculated that the species was na- JERDON (1851) first described M. destructor, reporting tive to North Africa, having spread westward from Egypt. it was "common in all parts of India." WROUGHTON (1892) Although WILSON (1962) wrote that M. destructor probably wrote that M. destructor was common in houses in South originated in tropical Asia, WILSON & TAYLOR (1967) la- India, but in northern India "above the Ghâts, it is a wild ter concluded that M. destructor "probably originated, like species, but not common." Most subsequent authors have other members of the subgenus Parholcomyrmex, from Af-
Figs. 1 - 4: Monomorium destructor. (1) head of major worker; (2) lateral view of major worker; (3) dorsal view of major worker; (4) workers on peanut butter bait, Lautoka, Fiji (photos by E. Sarnat).
rica." DLUSSKY (1994) also concluded that M. destructor rear two-thirds or so of the gaster, which is dark brown to was African in origin. Following DLUSSKY (1994), WET- nearly black. Workers are polymorphic, showing a wide TERER & VARGO (2003) listed M. destructor as an African size range and distinct allometry, readily identified by the species. fine transverse striae on the vertex and deep metanotal When evaluating the native and exotic ranges of a spe- groove (BOLTON 1987, FERNÁNDEZ 2007). Junior synonyms cies, researchers may consider a spectrum of distributional, of M. destructor include Monomorium basale SMITH, historical, evolutionary, ecological, and genetic information 1858 from Sri Lanka (synonymized by FOREL 1894), Myr- (see WETTERER 2008). Evidence considered indicative of a mica ominosa GERSTÄCKER, 1859 and Myrmica atomaria species' native range includes older records largely con- GERSTÄCKER, 1859 both from Mozambique (both synony- fined to a single continuous geographic region, occurrence mized by DALLA TORRE 1893), Monomorium vexator in inland native communities, high genetic diversity, co- SMITH, 1861 from Indonesia (synonymized by DONIS- occurrence of species-specific symbionts, and proximity to THORPE 1942), and Myrmica gracillima from Syria or Pa- the ranges of closely related species. In contrast, evidence lestine – site not given (synonymized by BOLTON 1987). indicative of a species' exotic range includes the sudden Methods appearance and spread of the species through an area dis- continuous with other known populations, occurrence ex- To document the geographic distribution of M. destructor, clusively in coastal and highly disturbed environments, low I used both published and unpublished records. Some pub- genetic diversity due to a founder effect, absence of species- lished reports of M. destructor from Africa and the Ara- specific symbionts, and geographic isolation from closely bian Peninsula may be misidentifications of closely related related species. Monomorium species. I obtained unpublished site records from museum specimens in the collections of the American Taxonomy Museum of Natural History (AMNH), the Archbold Bio- Monomorium destructor workers (Figs. 1 - 4) are distinctly logical Station (ABS), the British Natural History Mu- bi-colored: light yellow to yellow-brown, except for the seum (BMNH), the Museum of Comparative Zoology
98
Fig. 5: Worldwide distribution of Monomorium destructor (some African and Middle Eastern records may be misidenti- fications of related species).
(MCZ), the National Museums Liverpool (NML), the Ox- tor came from 107 geographic areas (i.e., countries, island ford University Natural History Museum (ONHM), and groups, major Caribbean islands, and US states). These in- the Smithsonian Institute's National Museum of Natural clude many areas for which I found no previously pub- History (SI). In addition, I used on-line databases with col- lished records, e.g., Bahamas, Barbuda, Barbados, Bon- lection data of specimens in the Australian National In- aire, Comoro Islands, Curaçao, Grenada, Haiti, Honduras, sect Collection (ANIC) and the California Academy of Kenya, Nevis, Pakistan, Réunion, St. Lucia, St. Martin, St. Sciences (CAS). I also received unpublished records from Vincent, and Venezuela (Tabs. 1 - 7). Despite being rela- B. Taylor (Egypt), O. Paknia (Iran), J. Miles (Palau), J. Le- tively simple to distinguish, some M. destructor specimens wallen (Vietnam), B. Hoffmann (Australia), W. Haines in the MCZ had been previously identified as Monomorium (Hawaii), J. Fellowes (China), and M. Angela (Aruba). Fin- pharaonis (LINNAEUS, 1758), and it is possible that some ally, I collected M. destructor specimens on the Cape Ver- reports of M. pharaonis in the literature were misidenti- de islands, on numerous Caribbean Islands, and in Florida. fied M. destructor. I did not find any M. pharaonis speci- Geographic coordinates for collection sites came from mens in the MCZ misidentified as M. destructor. Because published references, from specimen labels, or I looked up M. pharaonis is much more widely-known as a pest than the coordinates using maps and geography web sites (e.g., M. destructor, it seems unlikely that many reports of M. earth.google.com, www.tageo.com, and fallingrain.com). destructor in the literature were actually misidentified M. For older references and specimens, many site names, par- pharaonis. One exception is the record of M. destructor at ticularly in Asia, are no longer in use or are now spelled Archbold Biological Station, Florida in EISNER & al. (1980), differently and I searched, not always successfully, to de- which was actually M. pharaonis (M. Deyrup, pers. comm.). termine current names. If a site record listed a geographic Few records of M. destructor come from temperate ar- region rather than a "point locale," and I had no other rec- eas (Fig. 5). The highest latitude records are from indoor ord for this region, I used the coordinates of the largest populations in England (London: one record from a clothes town within the region or, in the case of small islands and warehouse and another from a switch box, and Kew Gar- natural areas, the center of the region. Often, if one source dens; 51.5 °N; DONISTHORPE 1911) and one report from had many sites less than 10 - 20 km apart (e.g., ALFIERI Dansville, New York (42.6 °N; USBEPQ 1962) that indi- 1931, CLOUSE 2007, my collection sites in Cape Verde), I cated "it is doubtful, however, if the species will survive did not plot every site. I did not map records of M. destruc- out-of-doors in New York." RUZSKII (1923) reported the tor on boats or intercepted in transit by quarantine inspec- highest latitude outdoor record of M. destructor (as M. tors. I excluded several specimen records originally iden- gracillimum) from a sandy spit by the Caspian Sea in Chele- tified as M. destructor that BOLTON (1987) subsequently ken, Turkmenistan (39.4 °N). DLUSSKY & ZABELIN (1985) identified as Monomorium mayri FOREL, 1902. Undated spe- reported M. destructor (as M. gracillimum) from western cimens collected by Theodore Pergande in Honduras and Kopet Dag, Turkmenistan, and Qashqadaryo, Uzbekistan Martinique must have been collected before Pergande's (both 38 - 39 °N). PISARSKI (1967, 1969) listed M. destruc- death in 1916. A specimen from Mombassa, Kenya, iden- tor (as M. gracillimum) from five sites in Afghanistan from tified by Forel, was collected before Forel's death in 1931. 35.1 to 36.0 °N. The only other records of M. destructor at temperate latitudes > 35 ° come from urban areas: Mel- Results bourne, Australia (37.8 °S; CLARK 1941), Auckland, New I mapped specimen records of M. destructor from > 600 Zealand (36.9 °S; HEMBRY 2005), and two cities in Ten- sites worldwide (Fig. 5). Specimen records of M. destruc- nessee (Chattanooga: 35.1 °N; Memphis: 35.2 °N; see Tab.
99 Tab. 1: Earliest known records for Monomorium destruc- Tab. 2: Earliest known records for Monomorium destruc- tor from Asia and neighboring islands. Unpublished rec- tor from North Africa, the Middle East, and neighboring ords include collector, museum source, and site. BMNH = islands. Abbreviations as in Table 1. Some records from Natural History Museum in London, MCZ = Museum of this region may be misidentifications of closely related Comparative Zoology, SI = Smithsonian Institute. + = no species. known published record.
Earliest record Earliest record Palestine or Syria ≤ 1860 (SMITH 1861a; as M. gracillima) + India ≤ 1851 (JERDON 1851) Egypt ≤ 1862 (MAYR 1862; as M. gracillimum) + Sri Lanka ≤ 1858 (SMITH 1858; as M. basalis) Palestine/Israel ≤ 1863 (ROGER 1863; as M. gracillimum) + Philippines ≤ 1858 (MAYR 1865; as M. basale) Syria ≤ 1877 (MAYR 1877; as M. gracillimum) + Indonesia ≤ 1861 (SMITH 1861b; as M. vexator) Asian Turkey ≤ 1877 (MAYR 1877; as M. gracillimum) + Turkmenistan ≤ 1877 (MAYR 1877; as M. gracilli- ≤ mum) Eritrea ≤ 1877 (EMERY 1877; as M. basale)
+ Burma/Myanmar ≤ 1885-87 (EMERY 1889; as M. gra- Yemen ≤ 1880 (EMERY 1881; as M. gracillimum) ≤ cillimum) Algeria ≤ 1883 (ANDRÉ 1883; as M. gracillimum) + Singapore ≤ 1899 (Haviland, BMNH): no site Sudan ≤ 1884 (MAGRETTI 1884; as M. gracillimum) + Iran ≤ 1907 (RUZSKII 1907; as M. gracil- ≤ limum) Canary Islands ≤ 1892 (EMERY 1893b)
+ Vietnam ≤ 1907 (FOREL 1907; as M. gracilli-≤ Tunisia ≤ 1908 (EMERY 1908; as M. gracillimum) mum) Iraq ≤ 1917 (CRAWLEY 1920; as M. gracillimum) + Taiwan ≤ 1909 (WHEELER 1909) Libya ≤ 1931 (MENOZZI 1932; as M. gracillimum) + China ≤ 1927 (WHEELER 1927) Chad ≤ 1948 (BERNARD 1948) + Thailand ≤ 1928 (H. Hillman, SI): Bangkok Niger ≤ 1950 (BERNARD 1950; as M. gracillimum) + Christmas I. ≤ 1933 (DONISTHORPE 1935) Saudi Arabia ≤ 1975 (COLLINGWOOD 1985; as M. gra- + Japan ≤ 1948 (BERNARD 1948) ≤ cilimum)
+ Afghanistan ≤ 1948 (COLLINGWOOD 1961; as M. Cape Verde ≤ 1983 (M.L. Lobo Lima, BMNH): Mindelo ≤ gracillimum) Lebanon ≤ 1988 (KUGLER 1988; as M. gracillimum) + Papua New Guinea ≤ 1957 (WILSON & TAYLOR 1967) Kuwait ≤ 1988-89 (COLLINGWOOD & AGOSTI 1996) + Nepal ≤ 1962 (R.L. Coe, BMNH): Taplejung United Arab ≤ 1995 (COLLINGWOOD & al. 1997) + Bangladesh ≤ 1967 (ALAM 1967 in HANNAN 2003) Emirates
+ Uzbekistan ≤ 1985 (DLUSSKY & ZABELIN 1985; Cyprus ≤ 2000 (CAB 2000 in DAFF 2001) ≤ as M. gracillimum)
+ Hainan Island ≤ 1995 (WU & WANG 1995) Santo Antão, São Nicolau, Fogo, and Brava. In the West Indies, I found M. destructor at scattered sites on most is- + Malaysia ≤ 1997 (WAY & BOLTON 1997) lands visited, usually in urban areas. My specimens were the + Pakistan ≤ 2007 (S & Z Valliani, MCZ): first records of M. destructor for several islands (Tab. 5). ≤Karachi Although M. destructor workers readily tend a wide range of Hemiptera and Lepidoptera (e.g., WROUGHTON 1892, DEYRUP & al. 2000, FIEDLER 2006), I found no rec- 7). RUZSKII (1907) reported M. destructor (as M. gracilli- ords of any species-specific symbionts associated with M. mum) from Southern Europe, but this location was too gen- destructor. eral for me to place a point on the map. ESPADALER (2005) Related Monomorium species reported M. destructor intercepted and exterminated in Bar- celona, Spain (41.3 °N) in a shipment from Busan, South In addition to M. destructor, BOLTON (1987) placed five Korea (35.1 °N), but gave no indication that M. destruc- other species in the Monomorium destructor species-group: tor was established at either site, so I did not map them. M. mayri, M. oscaris FOREL, 1894, M. robustior FOREL, In Cape Verde, I collected M. destructor in urban areas 1892, M. emeryi MAYR, 1895, and M. epinotale SANT- on six of the nine inhabited islands: Santiago, São Vicente, SCHI, 1923. Four members of the group (M. oscaris, M. ro-
100 Tab. 3: Earliest known records for Monomorium destruc- Tab. 4: Earliest known records for Monomorium destruc- tor from sub-Saharan Africa and neighboring islands. Ab- tor from Australia and Pacific Ocean islands. Abbrevia- breviations as in Table 1, and CAS = California Academy tions as in Table 1. of Sciences, MP = Museum Paris, ZSM = Zoologische
Staatssammlung München. Some records from this re- gion may be misidentifications of closely related species. Earliest record Hawaii ≤ 1896-97 (EMERY 1899)
Earliest record Australia ≤ 1910 (FOREL 1910) + Mozambique ≤ 1858 (GERSTÄCKER 1859; as M. Mariana Islands ≤ 1911 (WHEELER 1912) ≤ ominosa & M. atomaria) Society Islands ≤ 1925 (WHEELER 1936) + South Africa ≤ 1862 (MAYR 1862; as M. gracilli-≤ ≤ mum) Cook Islands ≤ 1925 (WILSON & TAYLOR 1967)
+ Madagascar ≤ 1899 (Grandidier, MP): site unknown FS Micronesia ≤ 1937 (CLOUSE 2007)
ANTSCHI + Guinea ≤ 1914 (S 1914; as M. gra- Marshall Islands ≤ 1947 (COLE 1949) ≤ cillimum) Solomon Islands ≤ 1953 (WILSON 1962) + Kenya ≤ 1931 (Prager, ZSM): Mombassa Gilbert Islands ≤ 1956 (E.S. Brown, BMNH): Tarawa + Central African Rep. ≤ 1948 (WEBER 1964) Wake Island ≤ 1957 (WILSON & TAYLOR 1967) + Congo (Zaire) ≤ 1948 (WEBER 1964) Samoa ≤ 1962 (WILSON & TAYLOR 1967) + Mauritius ≤ 1953 (MAMET 1954) Tuamotu Islands ≤ 1996 (MORRISON 1997) + Seychelles ≤ 1975 (MÜHLENBERG & al. 1977) Niue ≤ 1996 (COLLINGWOOD & VAN HARTEN + Cameroon ≤ 2006 (KENNE & al. 2006) ≤ 2001)
+ Réunion ≤ 2007 (B.L. Fisher, CAS): Le Port Galapagos ≤ 1997 (PEZZATTI & al. 1998)
+ Comoro Islands ≤ 2007 (B.L. Fisher, CAS): Mayotte; Fiji ≤ 2004 (WARD & WETTERER 2006) ≤ Tanaraki New Zealand ≤ 2004-5 (HEMBRY 2005) bustior, M. emeryi and M. epinotale) are known only from Palau ≤ 2005 (OLSEN & MILES 2005) sub-Saharan Africa (BOLTON 1987). BOLTON (1987) record- Phoenix Islands ≤ 2006 (PIERCE & al. 2006) ed M. mayri from Africa, the Arabian Peninsula, and Asia. Morphologically, M. destructor most closely resem- bles M. mayri, M. oscaris, and M. robustior, though the (1987) re-identified as M. mayri. It seems likely that other latter three species are uniform in color, not bi-colored like records of M. destructor may also be misidentified M. M. destructor (BOLTON 1987). BOLTON (1987) wrote that M. mayri (if this is, in fact, a valid species), particularly spe- mayri matched "the description of destructor in all respects cimens originally identified as M. gracillimum, a slightly except colour, mayri being uniformly dark brown, some- darker variant of M. destructor. times with a paler patch at the base of the first gastral ter- Many questions persist concerning the taxonomy of M. gite. I have decided to retain mayri as a valid species, sep- destructor and its close relatives. BOLTON (1987) wrote: "it arate from destructor, for the time being. The colour char- is possible that the names ominosa and atomaria, both des- acter is admittedly feeble but appears to be consistent." cribed from East Africa and subsequently synonymized BOLTON (1987) concluded: "the closest relative of oscaris with destructor, may represent early records of [Monomo- appears to be the pantropical tramp-species destructor, but rium] oscaris. However, as the original descriptions are so the two are separable by the shape of the petiole node in poor, and as the specimens involved seemingly have long dorsal view, especially in larger workers. In destructor the since disappeared, there is no way of proving this; in con- node is globular to subglobular but in oscaris it is strongly sequence they are left undisturbed as junior synonyms of anteroposteriorly compressed and markedly transverse. Also, destructor." Although I believe it is very likely that M. omi- at any given worker size, the scapes tend to be longer in nosa and M. atomaria are not M. destructor, but rather one destructor than oscaris." BOLTON (1987) reported: "the of its close relatives that are much more common in sub- colour and habitus of robustior approaches that of mayri Saharan Africa, I have followed BOLTON (1987) and in- most closely." However, several other characters of M. cluded these records. robustior differ from those of M. destructor and M. mayri: In Nigeria and Ghana, ROOM (1971), MAJER (1976), M. robustior has a narrower worker size range, larger eyes, and TAYLOR (1981) reported a common forest ant species and longer hairs on the first gastral tergite (BOLTON 1987). that they designated Monomorium sp. G. BOLTON (1987) Several specimens identified as M. destructor by earlier subsequently assigned Monomorium sp. G to M. oscaris, authors (e.g., from Egypt; DONISTHORPE 1942), BOLTON with the caveat that ants of this West African population
101 Tab. 5: Earliest known records for Monomorium destruc- Tab. 6: Earliest known records for Monomorium destruc- tor from the West Indies. Abbreviations as in Table 1, tor from South and Central America. Abbreviations as in and AMNH = American Museum of Natural History. Table 1.
Earliest record Earliest record
+ Jamaica ≤ 1893 (ANDRE 1893; as M. basale) + Brazil ≤ 1895 (FOREL 1895)
+ Puerto Rico ≤ 1908 (SMITH 1936) + Honduras ≤ 1916 (T. Pergande, SI): site unknown
+ Martinique ≤ 1916 (T. Pergande, SI): site unknown + Venezuela ≤ 1930 (J.G. Myers, MCZ): San Fernando ≤ de Apure + Cuba ≤ 1918 (W.M. Mann, SI): El Caney + Ecuador ≤ 1960 (KEMPF 1960) + Antigua ≤ 1918 (WHEELER 1923) + Colombia ≤ 1972 (KEMPF 1972) + Haiti ≤ 1926 (C.N. Wolcott, SI): Port au Prince
+ Dominican ≤ 1929 (MENOZZI & RUSSO 1930) Tab. 7: Earliest known records for Monomorium destruc- + Republic tor from the United States and Europe. Abbreviations as in Table 1. + Virgin Islands ≤ 1937 (BEATTY 1944)
+ Trinidad ≤ 1948 (BMNH): site unknown Earliest record
+ Nevis ≤ 1967 (N.L.H. Krauss, SI): site unknown Alabama ≤ 1905 (WHEELER 1906)
+ Montserrat ≤ 1967 (N.L.H. Krauss, SI): Plymouth Florida ≤ 1905 (WHEELER 1906)
+ Bahamas ≤ no date (N. Krauss, AMNH): Nassau Southern Europe ≤ 1907 (RUZSKII 1907; as M. gracillimum)
+ Barbuda ≤ no date (Jefferys, BMNH): site unknown England ≤ 1910 (DONISTHORPE 1911)
+ Guadeloupe ≤ 1986-87 (JAFFE & al. 1991) Tennessee ≤ 1932 (L.C. Murphree, SI): Memphis & ≤ Chattanooga + Bonaire ≤ 1999 (E. Scholtens & B. Krause, MCZ): ≤ Kaya Otomac New York ≤ 1961 (USBEPQ 1962 + Barbados ≤ 2003 (J.K. Wetterer, MCZ): Black Rock (SMITH, 1858). Monomorium destructor superficially re- + Grenada ≤ 2003 (J.K. Wetterer, MCZ): St. George's sembles the widespread pharaoh ant, M. pharaonis. Mono- + Tobago ≤ 2003 (J.K. Wetterer, MCZ): Bon Accord morium pharaonis workers vary in color, even within a single colony, from uniform yellow to yellow with a dark + St Lucia ≤ 2003 (J.K. Wetterer, MCZ): Pointe Hardy brown rear of the gaster. Monomorium destructor, how- ever, can be easily distinguished from M. pharaonis be- + St Vincent ≤ 2004 (J.K. Wetterer, MCZ): Calliaqua cause the head, alitrunk and pedicel of the workers are al- most entirely smooth and shiny in M. destructor, but are + Curaçao ≤ 2004 (J.K. Wetterer, MCZ): Ascension matte (not shiny) in M. pharaonis. In addition, M. destruc- + St Kitts ≤ 2007 (J.K. Wetterer, MCZ): Major's Bay tor has a much broader range of worker size than does M. pharaonis (M. destructor: 1.8 - 3.5 mm length; M. phara- + Aruba ≤ 2007 (J.K. Wetterer, MCZ): Palm Beach onis: 2.2 - 2.4 mm length; BOLTON 1987).
Discussion "may eventually prove to be separable at the species-level Distribution and impact from the eastern and southern population" of M. oscaris. Monomorium destructor has a widespread distribution in Monomorium sp. G, however, is distinctly bi-colored (TAY- tropical and subtropical parts of the Old World, except in LOR 2007). Although Monomorium sp. G differs from M. sub-Saharan Africa. In the New World, however, it is destructor in having only a narrow range of small wor- widespread only in Florida, the West Indies, and the Gala- kers with a different allometry, TAYLOR (2007) provision- pagos Islands (Fig. 5). ally regarded them as conspecific with M. destructor. Here, In the past, M. destructor was primarily spread by ship. however, I have been conservative and followed BOLTON For example, CLARKE (1922) reported M. destructor was a (1987) in considering Monomorium sp. G as distinct from serious problem on steamers traveling between California M. destructor. and the East Coast US via Panama Canal, writing that M. In addition to M. destructor, several other Monomorium destructor "not only caused a considerable pecuniary loss are considered major tramp species, e.g., Monomorium flo- in the destruction of food stuffs but attacked passengers and ricola (JERDON, 1851), M. monomorium BOLTON, 1987, M. crew… They would find their way in small or large num- pharaonis, M. sechellense EMERY, 1894, and M. subopacum bers into the beds and their bites were very painful." WEBER
102 (1939) found live M. destructor in his luggage in Massa- Remarkably, no subsequent collector found M. destructor chusetts several days after returning from Cuba by ship (not on Loggerhead Key (WETTERER & O'HARA 2002). Else- mapped). Now, air travel, combined with the ant's pro- where in the Florida Keys, DEYRUP (1991) wrote that M. pensity of nesting in electrical and electronic equipment, destructor "is spectacularly common on Key West." In a allows the possible spread of M. destructor to virtually any- recent visit to Key West, however, I was unable to find any where in the world. For example, ENGST (2005) reported M. destructor. Instead, all areas where I collected on Key that, on arrival in New Zealand, an air passenger found M. West were dominated by P. megacephala and/or S. invicta destructor living inside a sealed iPod bought in the air- (J.K. Wetterer, unpubl.). port in Fiji. Monomorium destructor populations appear to be ex- I found little information concerning possible impacts panding in the West Indies (see Tab. 5), where they are a of M. destructor in natural ecosystems, though there is on- serious household problem. For example, in my apartment going research in the Galapagos Islands, where expand- in Tunapuna, Trinidad, enormous trails of M. destructor ing M. destructor populations have caused concern (e.g., foragers streamed out of an electrical socket and into the VON AESCH & CHERIX 2005). cupboards, trashcan, and kitchen sink, carrying off any scrap In contrast, I found many reports of M. destructor des- of food they could find. In an electronics store in Aripo, troying property and attacking people. For example, STONEY Trinidad, the owners reported this species nesting inside (1995) wrote that in Western Australia, M. destructor "has their computers. In a hotel room in Grand Anse, Grenada, been known to chomp through a grown man's thong ov- a large trail of M. destructor emerged out of the air-condi- ernight and chew up anything from polystyrene cups to tioner to forage in my trashcan. In addition to urban areas, wiring in cars, telephones and houses... The insect has even I also found M. destructor swarming down trees in parks nibbled on newborn babies sleeping in their cots and left and disturbed forests, for example in beachfront parks on big holes in car tyres. 'Kids are virtually being eaten alive Grenada, St. Croix, St. Vincent, and St. Lucia and in a while they sleep at night,' the Derby shire president, Mr. forest of the poisonous manchineel tree (Hippomane man- Peter McCumstie, said." CHIN (1998) wrote how in Darwin, cinella LINNAEUS, 1753) on Curaçao. Australia, M. destructor "can cause havoc in the house- Native and exotic range hold since they bite and may occur almost everywhere in- side the house, feeding on a wide variety of food materi- There is consensus that M. destructor originated in the Old als. They frequently nest in power sockets and chew on World, but disagreement as to where in the Old World it electrical wiring and in some cases have started electrical is native (see Introduction). Because M. destructor was al- fires." In the community of Nguiu on Bathurst Island, Aus- ready known from many sites in both Asia and Africa in tralia, B. Hoffmann investigated an enormous outbreak of the 19th century (Tabs. 1 - 3), when the world's ant fauna M. destructor (CSIRO 2003). Hoffmann reported, "The was still very poorly documented, determining its native magnitude of damage is really overwhelming… There were range and reconstructing its early spread as an exotic is not massive trails going into houses from all directions – milli- possible using specimen records alone. ons and millions of Singapore ants [M. destructor] swarm- The close resemblance between M. destructor and sev- ing everywhere. These ants get into power points, they eat eral African species (BOLTON 1987) supports the proposal electrical wiring and short-circuit the power – two houses that M. destructor originated in Africa. However, DLUSSKY have already burned down recently because of damage & ZABELIN (1985) wrote that the distribution of its clos- caused to electrical systems'' (ANONYMOUS 2003). On Tobi est (unspecified) relatives suggested that M. destructor (as Island and Helen Reef Atoll in Palau, BOUDJELAS (2006) M. gracillimum) originated in Indo-Malaysia. In fact, M. reported that M. destructor was a serious threat to essen- destructor shows a fairly continuous distribution from North tial infrastructure and "causes extensive economic damage Africa, across the Middle East and Central Asia, all the way in human settlements by damaging fabric and rubber goods to Southeast Asia. Throughout this region, it occurs in di- and removing insulation from electric cables." LEE & al. verse natural and disturbed habitats. For example, COLLING- (2002) found that in surveys of food preparation outlets on WOOD & AGOSTI (1996) wrote that M. destructor "abounds Penang Island, Malaysia, M. destructor was the dominant ant throughout Arabia and seems to occur in every type of hab- species, making up 27.8% of the ant specimens collected. itat with many collections from Saudi Arabia, Oman and In the Dry Tortugas, the outermost of the Florida Keys, Yemen." MAYOR (1922) reported that M. destructor on Loggerhead There seems to be little rationale for considering M. de- Key was "a great pest in the wooden buildings of Tortu- structor native to Asia, but exotic to North Africa. Part of gas Laboratory, making its nests in crevices of the wood- the explanation for past researchers considering only Asia work. So voracious are these insects that we are obliged to as the original native range of M. destructor may be that swing our beds from the rafters and to paint the ropes with many M. destructor specimens, particularly from North a solution of corrosive sublimate, while all tables must have Africa and Arabia, were originally identified as M. gra- tape soaked in corrosive sublimate wrapped around their cillimum (e.g., see Tab. 2). Thus, before BOLTON (1987) legs if ants are to be excluded from them. These pests have designated M. gracillimum to be a junior synonym of M. the habit of biting out small pieces of skin, and I have seen destructor, most known Old World records of M. destruc- them kill within 24 hours rats which were confined in cages." tor came from Asia. At Tortugas Laboratory, "one of the scientists, a newcomer Records of M. destructor from sub-Saharan Africa are who allowed his sheet to touch the floor, was stung so badly rare and some of these may be misidentifications (Fig. 5; by a swarm of them that he lapsed into unconsciousness see Results). HETERICK (2006) considered M. destructor for a while" (STEPHENS & CALDER 2006). The Tortugas to be introduced to Madagascar. It seems likely that M. de- Laboratory buildings were later abandoned and torn down. structor is not native to sub-Saharan Africa as well. It also
103 seems likely that M. destructor is exotic to Australia and East, Pakistan, India, the Galapagos, and the Florida Keys, Oceania (WILSON & TAYLOR 1967). Whether or not M. but is also a house pest in more humid regions. Adapta- destructor is native to East and Southeast Asia (including tion to arid habitats may preadapt M. destructor to thrive Singapore) is an open question. Future research on the gen- in dry indoor conditions. Although M. destructor is not etic diversity of M. destructor populations in different parts yet known from many tropical and subtropical areas (Fig. of the world and the phylogeny of Monomorium should 5), I expect that it will continue to spread, particularly in help elucidate further the native and exotic ranges of M. arid areas such the Pacific coast of Central and South Am- destructor. erica and much of Mexico. The disappearance of M. destructor on Loggerhead Key Common names (see above) suggests that outbreaks may be relatively short- Although most authors have used no common name for M. lived. DEYRUP & al. (1988) wrote that M. destructor may destructor, some reports have used the name "Singapore be on the decline in the Florida Keys, and ESPADALER & ant" (e.g., HOFFMANN & O'CONNOR 2004, BOUDJELAS BERNAL (2003) noted that M. destructor had not been re- 2006; however, CHIN (1998) used this name for both M. corded in the Canary Islands for many years. A similar de- destructor and M. floricola). I could find no notable con- cline might be expected in areas currently experiencing nection between M. destructor and Singapore to explain this great outbreaks of M. destructor, including Cape Verde, common name. In the earliest use of "Singapore ant" that Bathurst Island, and Tobi Island. This pattern of popula- I found, BASEDOW (1918) wrote of "the so-called 'Singa- tion explosion followed by decline should be taken into con- pore ant,' whose tiny dimensions were by no means in pro- sideration in any large-scale efforts to control these ants. portion to their voracity." Unfortunately, BASEDOW (1918) Future research on the phylogeny of Monomorium, and did not include the Latin name for this species. In the ear- on the genetic diversity of M. destructor populations in dif- liest association I found between a particular ant species and ferent parts of the world should help elucidate further the the name "Singapore ant," CLARK (1938) wrote of "Mono- native and exotic ranges of this species. morium pharaonis, the well known house pest, a minute yellow ant, commonly known in Australia as the Singa- Acknowledgements pore Ant." I thank A. Wetterer, M. Wetterer, B. Taylor, J. Miles, and The name Singapore ant may have become associated B. Hoffmann for comments on this manuscript; S. Cover with M. destructor due to the mistaken synonymy of Myr- for help, encouragement, and ant identification; B. Bolton mica vastator SMITH, 1857 (= M. pharaonis), which was (BMNH), J. Carpenter (AMNH), S. Cover (MCZ), M. Dey- described from Singapore. WROUGHTON (1892) consider- rup (ABS), J. Hogan (ONHM), G. Knight (NML), and T. ed M. vastator a synonym of M. basale (= M. destructor) Schultz (SI) for help with their respective ant collections; and DALLA TORRE (1893) synonymized M. vastator with B. Taylor, O. Paknia, J. Miles, J. Lewallen, B. Hoffmann, M. destructor, apparently based on mislabeled specimens W. Haines, J. Fellowes, and M. Angela for providing un- in the British Museum. DONISTHORPE (1932), however, con- published records; W. O'Brien for GIS help; C. Ely and E. cluded "it is clear that the type of M. vastator Smith is at Belogay for translating Russian journal articles; D.P. Woj- Oxford, and that the species is really M. pharaonis L." cik and S.D. Porter for compiling their invaluable FORMIS Although FOREL (1911) reported M. destructor mayri bibliography; C. Scheid and R. Pasos of the FAU library (= M. mayri) from Singapore, the earliest publication I for processing so many interlibrary loans; E. Sarnat for use found reporting true M. destructor from Singapore was of his excellent photos; Florida Atlantic University, the Na- BOLTON (1987), who listed the collection site as Sabang, tional Science Foundation, and the National Geographic a locale I could not confirm (though there are eight towns Society for financial support. named Sabang in neighboring Indonesia and one in Ma- laysia). I did, however, find museum specimens of M. de- Zusammenfassung structor from Singapore (collected in 1899 and 1933; e.g., Die Ameisenart Monomorium destructor (JERDON, 1851), Tab. 1). auch "destroyer ant" genannt, ist ein Lästling in vielen tro- The common name "ninja ant," used in Cape Verde for pischen und subtropischen Gebieten, wo sie für das Abna- M. destructor, comes from the Ninja of feudal Japan, spies gen der Isolation von elektrischen Leitungen, das Behausen and assassins trained in martial arts, and popularized in und Zerstören von Elektrogeräten und das Attackieren von many recent movies and cartoons. This name, no doubt, re- Menschen berüchtigt ist. Ich habe veröffentlichte und un- lates to these ants' vicious attacks on people. Unfortunate- veröffentlichte Nachweise von > 600 Fundorten weltweit ly, this common name, like "Singapore ant," implies a spe- verortet, um die Ausbreitung von M. destructor zu evalu- cific Asian origin of this species, which is far from certain. ieren. Ich habe die frühesten bekannten Nachweise von M. DEYRUP & al. (2000) proposed a new common name destructor für 107 geographische Gebiete (Länder, Insel- for M. destructor: the "destructive trailing ant." Here, I pro- gruppen, große karibische Inseln und US-Bundesstaaten) pose a simpler, yet distinctive common name, based direct- zusammengetragen, einschließlich vieler, für die ich keine ly on its Latin name: the "destroyer ant." The equivalent bisher veröffentlichten Nachweise gefunden habe: Baha- Spanish common name would include its specific Latin mas, Barbados, Barbuda, Bonaire, Curaçao, Grenada, Haiti, name: "hormiga destructor." Honduras, Kenia, Komoren, Nevis, Pakistan, Réunion, St. Lucia, St. Martin, St. Vincent, Tobago und Venezuela. Mo- Future trends nomorium destructor hat die größte morphologische Ähn- Monomorium destructor appears to be most common as a lichkeit mit mehreren afrikanischen Monomorium-Arten serious pest in disturbed arid habitats in the tropics and sub- und eine offenbar durchgehende Verbreitung von Nord- tropics, including Cape Verde, North Africa, the Middle afrika bis Südostasien. Diese Umstände weisen auf Nord-
104 afrika als Ursprung von Monomorium destructor hin, aber CLOUSE, R.M. 2007: The ants of Micronesia (Hymenoptera: For- machen auch wahrscheinlich, dass die Art auch im Gebiet micidae). – Micronesica 39: 171-295. von Mittel-Ostasien bis Südasien nativ ist. Monomorium COLE, A.C. 1949: The ants of Bikini Atoll, Marshall Islands. – destructor tritt als Lästling am häufigsten in menschlich PanPacific Entomologist 25: 172-174. überprägten ariden bis semi-ariden Lebensräumen der Tro- COLLINGWOOD, C.A. 1961: The third Danish expedition to Cen- pen und Subtropen auf. Vorkommen von M. destructor tral Asia. Zoological Results. 27. Formicidae (Insecta) from scheinen oft lokal begrenzt und kurzlebig zu sein. Dieses Afghanistan. – Videnskabelige Meddelelser fra Dansk Natur- Muster einer explosiven Populationsentwicklung gefolgt historisk Forening 123: 51-79. von einer Abnahme der Population sollte bei großräumigen COLLINGWOOD, C.A. 1962: Some ants (Hym. Formicidae) from Bekämpfungsmaßnahmen gegen die Art berücksichtigt north-east Asia. – Entomologisk Tidskrift 83: 215-230. werden. COLLINGWOOD, C.A. 1985: Hymenoptera: Fam. Formicidae of Saudi Arabia. – Fauna of Saudi Arabia 7: 230-302. References COLLINGWOOD, C.A. & AGOSTI, D. 1996: Formicidae (Insecta: ALFIERI, A. 1931: Contribution à l'étude de la faune myrméco- Hymenoptera) of Saudi Arabia (Part 2). – Fauna of Saudi Ara- logique de l'Egypte. – Bulletin de la Société Entomologique bia 15: 300-385. d'Egypte 15: 42-48. COLLINGWOOD, C.A., TIGAR, B.J. & AGOSTI, D. 1997: Introduced ALLEN, C.R., EPPERSON, D.M. & GARMESTANI, A.S. 2004: Red ants in the United Arab Emirates. – Journal of Arid Environ- imported fire ant impacts on wildlife: A decade of research. – ments 37: 505-512. American Midland Naturalist 152: 88-103. COLLINGWOOD, C.A., POHL, H., GÜSTEN, R., WRANIK, W. & VAN ANDRE, E. 1883: Les fourmis [part]. – Species des Hyménoptères HARTEN, A. 2004: The ants (Insecta: Hymenoptera: Formici- d'Europe et d'Algérie 2: 281-344. dae) of the Socotra Archipelago. – Fauna of Saudi Arabia 20: 473-495. ANDRE, E. 1893: Description de quatre espèces nouvelles de four- mis d'Amérique. – Revue d'Entomologie (Caen) 12: 148-152. COLLINGWOOD, C.A. & VAN HARTEN, A. 2001: The ants (Hym., Formicidae) of Niue, south west Pacific. – Entomologists' ANONYMOUS 2003: Foreign ants a disaster for Tiwi Islands. – Can- berra Times 27 December 2003. Monthly Magazine 137: 139-144. CRAWLEY, W.C. 1920: Ants from Mesopotamia and north-west BASEDOW, H. 1918: Narrative of an expedition of exploration in North-Western Australia. – Proceedings of the Royal Geo- Persia (concluded). – Entomologist's Record and Journal of graphical Society of Australasia. South Australian Branch 18: Variation 32: 177-179. 105-295. CSIRO 2003: Exotic ants threaten aboriginal communities. – CSIRO Media Release - Ref 2003/213. –
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